aqua International Journal of Ichthyology

Vol. 18 (1), 15 January 2012

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Cirrhilabrus nahackyi, a new wrasse (Perciformes; Labridae) from the South Pacific

Fenton Walsh1 and Hiroyuki Tanaka2

1) PO Box 389 Kuranda, Queensland 4881, Australia. Email: [email protected] 2) Jinguh Clinic, 2-2-79 Jinguh, Miyazaki, Miyazaki 880, Japan. E-mail: [email protected]

Received: 20 May 2011 – Accepted: 6 July 2011

Abstract erstreckt. Die Rückenflosse der neuen Art ist auch ein wenig Cirrhilabrus nahackyi, a new species of labrid fish found länger als bei C. bathyphilus. at Viti Levu, Fiji, and at Tongatapu in Tonga is described from five specimens, 30.9-65.0 mm SL, captured in 35-50 m depths on outer reef slopes. The new species closely Résumé resembles C. bathyphilus from the Coral Sea. However, ter- Cirrhilabrus nahackyi, une nouvelle espèce de labre minal males differ in having an elevated pennant at the découverte à Viti Levu, Fidji et à Tongatapu, Tonga, est first and second dorsal spine. The male coloration of C. décrite sur base de cinq spécimens, de 30,9-65,0 mm SL, nahackyi also differs with regards to the dorsal and caudal capturée à une profondeur de 35-50 m sur des tombants fins. The spinous part of the dorsal fin is dusky yellow and récifaux du large. La nouvelle espèce ressemble de près à C. lacks a violet band, while the soft portion of this fin has a bathyphilus de la Mer de Corail. Néanmoins, les mâles distinctive yellow base, and larger red mid-dorsal band, adultes se distinguent par une haute excroissance sur la which is not present in C. bathyphilus. Additionally, the première et la deuxième épine dorsale. La coloration du caudal fin of the new species is red with only a thin sub- mâle de C. nahackyi diffère aussi pour ce qui concerne les marginal black line in the upper half of the fin in terminal nageoires dorsale et caudale. La partie dure de la dorsale est males compared to a yellow caudal fin in C. bathyphilus, d’un jaune sale et n’a pas de bande violette alors que la par- which has blue blotches on the membrane in the upper tie molle de cette nageoire a une base jaune nette et une part and has a broader black submarginal band extending plus large bande à mi-nageoire, ce qui n’existe pas chez C. the entire depth of the fin. The dorsal fin of the new bathyphilus. En outre, la caudale de la nouvelle espèce est species is also slightly longer than that of C. bathyphilus. rouge avec seulement une fine ligne noire submarginale dans la moitié supérieure de la nageoire chez les mâles Zusammenfassung adultes alors que la caudale de C. bathyphilus est jaune, Die neue Lippfisch-Art Cirrhilabrus nahackyi wird auf der avec des taches bleues sur la membrane dans la partie Grundlage von fünf Exemplaren mit 30,9-65,0 mm SL supérieure, et a une bande submarginale noire plus large beschrieben, die in 35-50 m Tiefe über äußeren Riffhängen qui s’étend sur toute la nageoire. La dorsale de la nouvelle bei Viti Levu, Fidschi-Inseln, und bei Tongatapu in Tonga espèce est aussi un peu plus longue que celle de C. bathy- gefangen wurden. Diese neue Art ähnelt stark C. bathyphilus philus. von den Korallenmeer-Inseln. Doch unterscheiden sich die ausgewachsenen Männchen durch einen aufragenden Wim- Sommario pel am ersten und zweiten Rückenflossenstrahl. Auch unter- Cirrhilabrus nahackyi, una nuova specie di labride rin- scheidet sich die Farbgebung der Männchen von C. nahackyi venuta a Viti Levu, Fiji e a Tongatapu a Tonga, è descritta an Rücken- und Schwanzflosse. Der dornartige Teil der sulla base di cinque esemplari di 30.9-65.0 mm SL, cat- Rückenflosse zeigt ein trübes Gelb, aber kein violettes Band; turati a 35-50 m di profondità sul versante esterno della der weiche Teil dieser Flosse hingegen weist eine deutlich barriera. La nuova specie è molto somigliante a C. bathy- gelbe Basis auf sowie ein größeres rotes Band im mittel-dor- philus del Mar dei Coralli. Tuttavia, i maschi terminali dif- salen Bereich, das bei C. bathyphilus fehlt. Außerdem ist die feriscono per avere un’estensione a guisa di bandierina sulla Schwanzflosse der neuen Art rot mit einer nur dünnen sub- prima e sulla seconda spina dorsale. Il maschio di C. marginalen schwarzen Linie in der oberen Hälfte bei aus- nahackyi differisce inoltre per la colorazione delle pinne gewachsenen Männchen; C. bathyphilus hingegen hat eine dorsale e caudale. La parte spinosa della dorsale è di color gelbe Schwanzflosse mit blauen Flecken auf der Membran giallo opaco e manca della banda violetta, mentre la parte im oberen Teil und einem breiteren schwarzen submar- molle ha una base gialla ben evidente e una banda rossa ginalen Band, das sich über die ganze Tiefe der Flosse mediana più larga che non è presente in C. bathyphilus. In

1 aqua vol. 18 no. 1 - 15 January 2012 Cirrhilabrus nahackyi, a new wrasse (Perciformes; Labridae) from the South Pacific aggiunta, la pinna caudale della nuova specie è rossa con John Randall from the Bishop Museum in Hawaii una sottile linea nera submarginale nella metà superiore informed the first author that Bruce Carlson from della pinna rispetto ad una pinna caudale gialla in C. the Georgia Aquarium (Atlanta, USA) had received bathyphilus, che possiede macchie blu sulla membrana several fish from aquarium fish collector Tony della parte superiore e ha una banda submarginale nera più ampia che si estende per tutta l’altezza della pinna. La dor- Nahacky in Fiji and photographs confirmed it was sale della nuova specie è inoltre leggermente più lunga di probably conspecific with the undescribed Cirrhi- quella di C. bathyphilus. labrus from Tonga. Unfortunately the difficulty in removing this fish from a large public display meant that we were unable to examine it. The first author INTRODUCTION contacted the collector directly and our efforts to The labrid genus Cirrhilabrus Temminck & secure specimens were rewarded when Tony Schlegel, 1845 contains small, colorful and sexually Nahacky collected four fish and forwarded them to dimorphic coral-reef fishes that range across the the first author in December 2005. tropical and subtropical Indo-Pacific region. Prior to Cirrhilabrus bathyphilus was originally described 1958, only the following species were known: C. from the Coral Sea (Holmes Reef off north-eastern cyanopleura (Bleeker, 1851), C. temminckii Bleeker, Australia and Chesterfield Bank near New Caledo- 1853a, C. jordani Snyder, 1904, and C. exquisitus nia and now reported from Vanuatu) as the 42nd Smith, 1957. Four additional species, including C. member of the genus (Fig. 1). In the present paper solorensis Bleeker, 1853b, C. heterodon Bleeker, 1871, we describe a new species Cirrhilabrus nahackyi col- and C. lyukyuensis Ishikawa, 1904, have generally lected in Fiji and Tonga in the South Pacific as the been regarded as synonyms. However, some authors 48th species of Cirrhilabrus. now recognize C. solorensis as a valid species (Allen & Randall 1996; Allen & Kuiter 1999; Parenti & METHODS Randall 2000). In a recent paper describing C. beau- Lengths given for specimens are standard length perryi from Papua New Guinea, Allen et al. (2008) (SL), the straight-line distance from the median noted that the genus contains 46 valid species. Since anterior point of the upper lip to the base of the cau- then Randall and Tanaka (2009) added Cirrhilabrus dal fin (posterior end of the hypural plate). Head naokoae from Indonesia as the 47th species making length is measured from the median anterior point Cirrhilabrus the second most speciose genus in the of the upper lip to the posterior end of the opercu- family. At present only Halichoeres contains more lar membrane; snout length is from the same ante- species, Parenti & Randall (2011) noted 80 species, rior point to the fleshy edge of the orbit. Body depth although a recent molecular study has cast doubt on is the greatest depth measured to the base of the dor- the generic placement of several species presently sal spines and body width is the greatest width just included in Halichoeres (Barber et al. 2005). Reallo- posterior to the opercular flap. Orbit diameter is the cation of some of these species to other genera may greatest fleshy diameter and the interorbital width is result in Cirrhilabrus being the largest genus of the least bony width. Caudal peduncle length is Labridae. measured horizontally from the rear of the anal fin Allen et al. (2003) attributed many discoveries of to the base of the caudal fin and caudal peduncle new Cirrhilabrus over the past 30 years to the devel- depth is the least depth. Predorsal, pre-anal and pre- opment and availability of SCUBA equipment for pelvic lengths are taken from the upper lip to the scientific diving. Some of the more recently anterior origin of the respective fin. Lengths of each described species occur below 40-50 m. New tech- fin spine, ray and dorsal fin height are taken from nology involving mixed-gas and rebreather diving the base of each element. equipment has made these deeper habitats more Pectoral ray counts include the short rudimentary accessible to scientific surveys and further new dis- upper ray. The lateral line scale counts are given in coveries are likely. two parts, the anterior count from the upper end of The second author received an aquarium photo- the opercular flap to below the soft portion of the graph in 2002 from Larry Sharron in Tonga of a Cir- dorsal fin. The second or posterior lateral line count rhilabrus that appeared to be a close ally of C. bathy- is from the midlateral peduncular portion to the philus Randall & Nagareda, 2002. The fish was sold base of the caudal fin (a single scale usually located to aquarium fish dealers before the second author posterior to the base of the caudal fin is included). recognized it as a probable new species. In 2005 Gill raker counts include rudiments and only a total aqua vol. 18 no. 1 - 15 January 2012 2 Fenton Walsh and Hiroyuki Tanaka

Fig. 1. Holotype of Cirrhilabrus bathyphilus, AMS I.151103.001 male 48.7 mm SL, Holmes Reef, Coral Sea. Photo by F. Walsh. count is given as it is difficult to determine which width 2.0-2.2 in body depth; head length 2.7-3.1 in gill raker is at the angle. Type specimens are SL; snout length 3.85-4.15 in head length; pelvic fin deposited at the Queensland Museum, Brisbane short, not reaching base of anal fin, 4.3-5.2 in SL; (QM), United States National Museum, Washing- caudal fin rounded; eye large, orbit diameter 3.15- ton (USNM) and the University of Miyazaki- Fishes 3.85 in head length. Color in life of males: abdomen Science, Miyazaki (MUFS). pale yellow; body red dorsally fading to yellowish ventrally; soft portion of dorsal black outer band, Cirrhilabrus nachakyi, n. sp. red band mid-dorsally and broader posteriorly, yel- Nahacky’s Wrasse (Figs 2-5, Table I) low band at base; spinous portion black on the first membrane fading to irregular dusky on yellow ?Cirrhilabrus sp. Nahackyi’s Fairy Wrasse - Michael toward first soft ray, thin outer margin violet to pale 2009: 115-117. blue; nape dusky. Color in alcohol pale, males with ?Cirrhilabrus sp 1. Tongan Fairy Wrasse - Kuiter black submarginal band in dorsal fin, caudal fin 2010: 142. translucent. Largest specimen 65 mm SL. Description: Dorsal rays XI,9; anal rays III,9; first Holotype: QM I.38421, male, 55.6 mm SL, Bega dorsal and anal soft rays unbranched, all others Lagoon, Viti Levu, Fiji Islands, over gentle sloping branched, last to base; pectoral rays 15, upper two rubble bottom, 35 m, hand-net, December 2005. unbranched; pelvic rays I,5; principal caudal rays 13; Paratype: QM I.38242, female, 31.8 mm SL, same median 11 unbranched; upper and lower procurrent data as holotype; USNM 387558, male, 42.4 mm caudal rays 6, posteriormost segmented; lateral line SL, same data as holotype; MUFS 23365-23367, interrupted; dorso-anterior series of pored scales male, 52.8 mm SL, same data as holotype; QM 17+5, scales above lateral line to base of dorsal fin 2; I.38241, 65.0 mm SL, Tongatapu, Tonga, from the scales below lateral line to base of anal fin 6; median aquarium trade, June 2007. predorsal scales 5; median prepelvic scales 6; cir- Diagnosis: Dorsal rays XI,9; anal rays III,9; pec- cumpeduncular scales 16; horizontal scale rows on toral rays 15; lateral line scales 17+5; median pre- cheek 2; gill rakers14 (14-15); branchiostegal rays 5; dorsal scales 5; horizontal rows of scales on cheek 2; vertebrae 9 + 16. gill rakers 14-15; body depth 3.00-3.45 in SL; body Body depth 3.0 (3.10-3.45) in SL; body com-

3 aqua vol. 18 no. 1 - 15 January 2012 Cirrhilabrus nahackyi, a new wrasse (Perciformes; Labridae) from the South Pacific pressed, width 2.05 (2.0-2.2) in body depth; head located just below upper eye level and just forward to length 2.75 (2.7-3.1) in SL; dorsal profile of head front edge of eye; anterior nostril very short mem- convex; snout moderately pointed its length 4.05 branous tube, slightly higher posteriorly and located (3.85-4.15) in head length; orbit large 3.7 (3.15- anteroventral to posterior nostril, its diameter about 3.85) in head length; interorbital space convex, least equal to sensory pores of cephalic lateralis system. bony width 4.15 (3.85-4.35) in head length; caudal Suborbital pores from middle of eye to below front peduncle depth 2.3 (2.20-2.45); caudal peduncle edge of eye 12 (10-13); pores along free edge of pre- length 2.05 (2.05-2.40) in head. opercle 8 (7-8); pores on mandible to front of chin 4. Mouth terminal and oblique, forming angle of Scales cycloid; head scaled except interorbital approximately 30° to horizontal axis of body and space, snout and chin; opercle covered by seven large head; mouth small, maxilla extending just posterior scales; cheek with two horizontal rows of scales to vertical through anterior nostril, upper jaw length below eye; naked lower flange of preopercle thin, 4.4 (4.00-4.35) in head length; dentition typical of greatest width at angle about 2.5 in orbit diameter the genus, front and upper jaw with three pairs of in holotype; base of dorsal and anal fins with single canine teeth anteriorly at side of upper jaw, anterior row of large elongated scales, one per membrane; pair forward projecting, next two pairs increasing in last pored scale on lateral line at base of caudal fin length, more recurved and laterally projecting; upper enlarged and pointed; terminal scale on midline just with closely set small conical teeth (16 in holotype posterior to last pored scale greatly enlarged and posterior to third canine); lower jaw with single pair pointed; no scales on paired fins; pelvic fins with of forward and laterally projecting canines and median ventral process of two elongate scales about closely set small conical teeth, first five largest and three-fourths the length of pelvic spine, thin axillary just posterior of canines (18 in holotype); tongue scale of each pelvic fin about three-fourths the short and rounded. Gill rakers short, longest on first length of pelvic spine. gill arch and less than one-half of longest gill fila- Origin of dorsal fin above third lateral line scale; ments. predorsal distance 3.2 (3.00-3.35) in SL; dorsal fin Posterior margin of preopercle with 33 (28-34) height elevated in terminal males 1.75 (1.05-2.25) small serrae; edge of preopercle free from behind in head length (interspinous membranes of dorsal centre of eye to below anterior edge of pupil; lower fin extending above spine tips and supported by and rounded margin of preopercle thin and mem- slender fleshy rod behind first and second dorsal branous. spines, forming pennant); first dorsal spine 3.2 Posterior nostril subtriangular with short rim, (2.25-3.65) in head length, about equal to second

Fig. 2. Holotype of Cirrhilabrus nahackyi QM I.38421, male, 55.6 mm SL, Bega Lagoon. Fiji. Photo by F. Walsh. aqua vol. 18 no. 1 - 15 January 2012 4 Fenton Walsh and Hiroyuki Tanaka

Table I. Proportional measurements of type specimens of Cirrhilabrus nahackyi as percentage of standard length.

Holotype Paratypes

QM QM USNM MUFS QM I.38421 I.38242 387558 23365-67 I.38241

Sex male female male male male Standard length (mm) 55.6 31.8 42.4 52.8 65.0 Body depth 33.5 31.8 29.0 32.4 29.8 Body width 16.4 15.4 14.4 14.8 13.8 Head length 36.5 36.8 35.8 34.5 32.5 Snout length 9.0 9.1 9.2 8.3 8.5 Orbit diameter 9.9 11.6 9.9 9.1 8.6 Interorbital width 8.8 8.5 8.7 8.5 8.5 Upper jaw length 8.3 8.5 9.0 8.1 7.5 Caudal peduncle depth 15.8 15.7 14.6 14.6 14.6 Caudal peduncle length 17.8 17.6 15.3 14.4 16.0 Predorsal length 31.1 30.2 33.3 32.4 30.0 Preanal length 58.3 59.7 60.6 58.1 52.3 Prepelvic length 34.5 39.0 40.8 32.8 33.5 Dorsal fin base 63.3 61.6 59.2 58.1 61.7 Height of dorsal fin 20.7 16.4 18.9 21.2 30.6 First dorsal spine 11.3 10.1 10.1 12.9 14.5 Longest dorsal spine 14.4 15.7 15.1 16.7 16.3 Longest dorsal ray 18.5 16.4 18.9 18.2 17.2 Anal fin base 29.1 28.0 28.5 28.8 26.9 First anal spine 8.5 7.9 7.5 8.7 8.0 Second anal spine 9.9 11.6 9.7 10.4 9.5 Third anal spine 11.9 13.2 14.2 11.6 14.0 Longest anal ray 20.1 15.4 16.3 16.7 18.5 Caudal fin length 27.0 26.4 28.1 26.7 24.3 Caudal concavity 0 0 0 0 0 Pectoral fin length 23.9 23.6 24.8 22.7 23.1 Pelvic spine length 12.6 11.0 10.8 12.3 11.1 Pelvic fin length 23.4 21.4 19.6 19.3 20.8

Fig. 3. Paratype of Cirrhilabrus nahackyi QM I.38242, female, 31.8 mm SL, Bega Lagoon. Fiji. Photo by F. Walsh.

5 aqua vol. 18 no. 1 - 15 January 2012 Cirrhilabrus nahackyi, a new wrasse (Perciformes; Labridae) from the South Pacific spine in males; other dorsal spines subequal, the lowing centers of longitudinal scale series punctu- longest 2.55 (2.0-2.4) in head length; first or sec- ated by series of approximately eight irregularly ond soft dorsal ray longest 1.95 (1.90-2.25) in spaced violet spots on each line (stress coloration), head length; origin of anal fin vertically below last six faint violet-red longitudinal lines on head: three dorsal spine; preanal length 1.7 (1.65-1.90) in SL; above and three either side of eye extending from first anal spine 4.3 (3.95-4.75) in head; second anal snout to opercle; chest and lower third of head spine 3.7 (3.15-3.70) in head; third anal spine 3.1 below eye and posterior to chin abruptly pale yel- (2.3-3.0) in head; sixth, seventh or eighth anal soft lowish, shading to yellowish red on abdomen; nape rays longest, 1.8 (1.75-2.40) in head; caudal fin 3.7 dusky and extending to just below third dorsal (3.55-4.10) in SL, caudal fin rounded to slightly spine above lateral line; iris red-yellow; spinous rounded on males and slightly rounded on females; portion of dorsal fin yellow, first spinous mem- third pectoral ray longest 1.55 (1.40-1.55) in head; brane and pennant black, with broad dusky band pelvic fin short extending just beyond anus, longer centrally and shading to thin dusky yellow band at in males than females; second ray longest 1.55 first soft dorsal ray, base of each spine with dusky (1.55-1.85) in head, 4.3 (4.7-5.2) in SL. triangle; soft portion of fin distinctly differing Color of male holotype in alcohol: from spinous, with broad black submarginal band, pale yellowish, dusky on nape and extending to narrowing slightly posteriorly, broad red central below third dorsal spine; dorsal fin mostly dusky band broadening posteriorly, base of soft portion grey, first membrane black, soft portion with nar- similar to spinous, yellow with dusky triangle at row translucent outer margin and broad outer each dorsal ray base; anal fin yellow with blue-vio- black band narrowing posteriorly, translucent let margin and broad irregular red submarginal below and forming narrowing band running ante- band; caudal fin red shading to translucent distally, riorly to base of second dorsal ray; caudal fin faint line of dusky submarginal spots; pelvic fins translucent with few small dusky spots distally; translucent yellow, first two membranes yellow; remaining fins translucent. pectoral fins transparent, red to yellow at base. Color of the female paratypes in The largest male paratype from Tonga has an a l c o h o l : pale yellowish, nape dusky to below overall yellowish body and the four lines on the third dorsal spine; first dorsal fin membrane black; head are more discernible than in the holotype; all other fins translucent. and the caudal fin has a submarginal black line, Color of male holotype in life (Figs 2, thickest dorsally with a blue-violet margin. 4): bright red shading to yellowish red posteriorly, Color of female paratype in life (Fig. with several indistinct slightly darker red lines fol- 3): bright red body and head, shading to yellow on

Fig. 4. Paratype of Cirrhilabrus nahackyi QM I.38241, male, 65.0 mm SL, from Tongatapu. Tonga. Photo by H. Tanaka. aqua vol. 18 no. 1 - 15 January 2012 6 Fenton Walsh and Hiroyuki Tanaka abdomen and below eye, six violet lines with red snout to base of caudal fin, lower three lines on margin following centers of longitudinal scale either side of eye, lowest just below the horizontal series with irregular larger violet spots, uppermost mid-line; dorsal fin translucent yellow except for line extending from snout along mid-dorsal line black first dorsal spine; anal and caudal fins and dorsal fin base, other five lines extending from translucent yellow; paired fins transparent.

Fig. 5. Cirrhilabrus nahackyi, terminal male, 10cm TL, collected in 48 m from Tonga. Photo by L. Sharron.

Fig. 6. Cirrhilabrus cf. bathyphilus, QM I.38235 male, 69.4 mm SL, Efate Island Vanuatu. Photo by F. Walsh.

7 aqua vol. 18 no. 1 - 15 January 2012 Cirrhilabrus nahackyi, a new wrasse (Perciformes; Labridae) from the South Pacific

REMARKS REFERENCES The new species is known only from Fiji and ALLEN, G. R. 2000. Description of a new wrasse (Pisces: Tonga. It appears to be closely related to Cirrhilabrus Labridae: Cirrhilabrus) from Northern Sumatra, Indonesia. bathyphilus, which occurs in the Coral Sea off Aus- aqua, journal of Ichthyology and Aquatic Biology 4 (2): 45-50. ALLEN, G. R. & R. H. KUITER. 1999. Descriptions of two tralia, Vanuatu and New Caledonia. Both species new wrasses of the genus Cirrhilabrus (Labridae) from share several common morphological features such Indonesia. aqua, Journal of Ichthyology and Aquatic Biol- as general body coloration, size, and also are found ogy 3 (4): 133-140. in similar rubble habitats on outer reef slopes. The ALLEN, G. R. & J. E. RANDALL. 1996. Three new species of caudal fin shape, general coloration, and color of the wrasses (Labridae: Cirrhilabrus) from Papua New Guinea dorsal fin are key differences between the two and the Solomon Islands. Revue francaise d’Aquariologie species. Moreover, C. nahackyi has a rounded caudal 23 (3-4): 101-111. ALLEN, G. R., RANDALL, J. E. & CARLSON, B.A. 2003. Cir- fin in terminal males compared to the emarginate rhilabrus marjorie, a new wrasse (Pisces: Labridae) from shape of C. bathyphilus. The new species also lacks Fiji. aqua, Journal of Ichthyology and Aquatic Biology 7 the violet blue spots and thick submarginal black (3): 113-118. band on the caudal fin that are typical of C. bathy- BARBER, P. H. & BELLWOOD, D. R. 2005. Biodiversity philus. Instead, the caudal fin is pale red with only a hot spots: evolutionary origins of biodiversity in wrasses thin submarginal black band present in terminal (Halichoeres: Labridae) in the Indo-Pacific and new world males. The dorsal fin of the new species is dusky yel- tropics. Molecular and Phyogenetics and Evolution 35 low on the spinous portion, except for the first (2005): 235-253. KUITER, R. H. 2002. Fairy & rainbow wrasses and their rel- membrane, which is black and extended in height, atives. 208 pp. TMC Publishing, Chorleywood, UK. and the soft portion has three distinct bands of yel- KUITER, R. H. 2010. Labridae fishes: wrasses. 142 pp. low at the base, red centrally and a black outer band. Aquatic Photographics, Seaford, Australia. In contrast, C. bathyphilus has an outer black band MICHAEL, S. 2009. Wrasses and parrotfishes. The complete running the full length of the fin and a distinctive illustrated guide to their identification, behaviors, and cap- violet blue band in the middle of the dorsal fin. tive care. TFH Publications, Neptune City, New Jersey. It has come to our attention that another fish that PARENTI, P. & RANDALL, J. E. 2000. An annotated check- list of the species of the labroid fish families Labridae and is similar to C. nahackyi is found at Efate Island in Scaridae. J.L.B. Smith Institute of Ichthyology Ichthyologi- Vanuatu (Fig. 6). Larry Sharron supplied specimens, cal Bulletin 68: 1-97. which have become known as the “Hooded Fairy PARENTI, P. & RANDALL, J. E. 2011. Checklist of the Wrasse” in the aquarium trade as well as specimens species of the families Labridae and Scaridae: an update. from Tanna Island in southern Vanuatu. The latter Smithiana 13: 29-44. fish appears to be C. bathyphilus but the Efate Island RANDALL, J. E. 1992. A review of the labrid fishes of the labrid we believe is a subspecies of C. bathyphilus genus Cirrhilabrus from Japan, Taiwan and the Mariana Islands, with descriptions of two new species. Micronesica which we plan to describe in a future paper and will 25 (1): 99-121. illustrate the various colour forms of C. bathyphilus. RANDALL, J. E. & ALLEN, G. R. 2003. Paracheilinus rubri- caudalis, a New Species of Flasherwrasse (Perciformes: ACKNOWLEDGEMENTS Labridae) from Fiji. aqua, Journal of Ichthyology and We are especially grateful to Tony Nahacky who Aquatic Biology 7 (3): 103-112. supplied the four type specimens and provided RANDALL, J. E. & NAGAREDA, B. 2002. Cirrhilabrus bathy- detailed information on the type location for Cir- philus, a new deep-dwelling labrid fish from the Coral rhilabrus nahackyi. We are also very grateful to Sea. Cybium 26 (2): 123-127. RANDALL, J. E. & PYLE, R. L. 2001. Three new species of Larry Sharron for his initial recognition and infor- labrid fishes of the genus Cirrhilabrus from Islands of the mation of this new species and for supplying loca- Tropical Pacific. aqua, Journal of Ichthyology and Aquatic tion information for the fifth paratype specimen. Biology 4 (3): 89-98. We would also like to thank Dr John Randall for RANDALL, J. E. & KUITER, R. H. 1989. Cirrhilabrus punc- assisting greatly with his knowledge and his tatus, a new species of labrid fish from the southwestern encouragement, Dr Gerry R. Allen for his help in Pacific. Revue francaise d’Aquariologie 16 (2): 43-50. reviewing our manuscript, Bruce Carlson for some SENOU, H. & HIRATA, T. 2000. A new labrid fish, Cirrhi- labrus katoi, from southern Japan. Ichthyological Research of the first photographs the new species, Jeff John- 4 (1): 89-93. son of the Queensland Museum for assistance with SPRINGER, V. G. & RANDALL, J. E. 1974. Two new species the specimens and Sue Morrison of the Western of the labrid fish from the Red Sea. Israel Journal of Australian Museum for x-rays. Zoo logy 23: 45-54. aqua vol. 18 no. 1 - 15 January 2012 8 aqua, International Journal of Ichthyology

A new species of Dragonet (Synchiropus: Callionymidae) from Indonesia

Gerald R. Allen1 and Mark V. Erdmann2

1) Department of Aquatic Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, Perth, Western Australia 6986 2) Conservation International Indonesia Marine Program, Jl. Dr. Muwardi No. 17, Renon, Denpasar 80235 Indonesia

Received: 21 June 2011 – Accepted: 21 July 2011

Abstract Papouasie occidentale, Indonésie. L’espèce à également été Synchiropus tudorjonesi is described on the basis of 4 speci- signalée à Bali, Indonésie, à partir de photographies sous- mens, 21.1-35.5 mm SL, collected at Cenderawasih Bay, marines. Elle semble la plus proche de S. morrisoni, qui West Papua, in Indonesia. The species has also been occupe de grands espaces dans le Pacifique ouest. Les deux recorded from Bali, Indonesia, from underwater pho- espèces ont des distributions qui se chevauchent en tographs. It appears to be most closely related to S. mor- Indonésie, mais sont séparées écologiquement dans la risoni, which ranges widely in the western Pacific. The two mesure où S. tudorjonesi occupe des eaux plus profondes species have overlapping distributions in Indonesia, but are (50-70 m) que S. morrisoni qui évolue moins bas (12-33 m ecologically separated with S. tudorjonesi occurring in deeper environ). Les deux espèces ont des données méristiques et water (50-70 m) compared to the shallower depth range morphométriques similaires. Néanmoins, le mâle adulte (about 12-33 m) of S. morrisoni. Both species exhibit simi- de S. morrisoni a une première dorsale nettement plus lar meristic and morphometric features. However, the adult grande qui peut atteindre deux fois la hauteur de la sec- male of S. morrisoni has a much taller first dorsal fin, which onde dorsale (ou jusqu’à 2,5 en LS) alors que S. tudorjonesi may reach twice the height of the second dorsal fin (or to 2.5 a la nageoire bien plus courte (4,2 en LS). Il y a également in SL) compared to the much lower fin (4.2 in SL) of S. des différences notables de couleur, particulièrement eu tudorjonesi. There are also significant colour differences, par- égard à la première dorsale du mâle. ticularly with regards to the male’s first dorsal fin. Sommario Zusammenfassung Synchiropus tudorjonesi è descritto sulla base di 4 esem- Synchiropus tudorjonesi wird auf der Grundlage von vier plari di 21.1-35.5 mm SL, raccolti a Cenderawasih Bay, Exemplaren mit 21,1-35,5 mm SL beschrieben, die in der Papua Occidentale, Indonesia. La presenza della specie è Cenderawasih-Bucht, Westpapua, Indonesien gefangen stata segnalata anche a Bali, Indonesia, come documentato worden waren. Nachweise dieser Art durch Unterwasserfo- da fotografie subacquee. Appare molto vicina a S. morri - tografien gibt es auch von Bali, Indonesien. Am nächsten soni, diffusa ampiamente nel Pacifico occidentale. Le due verwandt ist offenbar die Art S. morissoni, die im west- specie hanno distribuzione simpatrica in Indonesia, ma lichen Pazifik weite Verbreitung gefunden hat. Die beiden sono ecologicamente separate, poiché S. tudorjonesi vive in Arten überschneiden sich zwar in der Verbreitung in acque più profonde (50-70 m) rispetto a quelle occupate Indonesien, sind aber ökologisch getrennt. S. tudorjonesi da S. morrisoni (circa 12-33 m). Entrambe le specie mo - lebt in tieferem Wasser (50-70 m), S. morrisoni im strano caratteristiche morfometriche e biometriche simili. flacheren (12-33 m). Die meristischen und morphome- Tuttavia, il maschio adulto di S. morrisoni ha una prima trischen Merkmale sind bei beiden Arten ähnlich. Doch pinna dorsale molto più elevata, che può raggiungere besitzt das Männchen von S. morrisoni eine deutlich altezze anche due volte superiori di quelle della seconda größere erste Rückenflosse, sie kann doppelt so hoch sein dorsale (oppure fino a 2.5 in SL), rispetto a S. tudorjonesi wie die zweite (bis 2,5 nach SL); bei S. tudorjonesi hinge- che possiede una pinna molto meno elevata (4.2 in SL). gen ist die erste Rückenflosse vergleichsweise niedrig (4,2 Inoltre, ci sono significative differenze di colorazione, ri- nach SL). Hinzu kommen deutliche Farbunterschiede, vor guardante in modo particolarmente la prima pinna dorsale allem bei der ersten Rückenflosse der Männchen. del maschio.

Résumé INTRODUCTION Synchiropus tudorjonesi est décrit sur base de 4 spécimens, Dragonets of the family Callionymidae are bot- 21,1-35,5 mm de LS, collectés à Cenderawasih Bay, tom-living fishes frequently found on sand, mud,

9 aqua vol. 18 no. 1 - 15 January 2012 A new species of Dragonet (Synchiropus: Callionymidae) from Indonesia or dead coral rubble. They usually rest on their Papua Province, Indonesia, 70 m, clove oil and outstretched pelvic fins and move about the reef in hand net, M. Erdmann, 20 September 2010. slow, short bursts. Males are generally more colour- Paratypes: MZB 20020, female, 21.2 mm SL, col- ful than females and use their ornate first dorsal fin lected with holotype; WAM P.33357-001, female, to signal their spawning intentions to nearby 24.4 mm SL, same collection data as holotype females. In some species the first dorsal is exceed- except 13 September 2010; WAM P.33368-002, ingly tall and sail-like and is fully erected during female, 25.2 mm SL, collected with holotype. courtship displays. More than 125 species are Diagnosis: A species of the subgenus Synchiropus known from the Indo-Pacific region. However, there (see Fricke 1981) with the following combination is disagreement concerning the generic classification of features: dorsal rays VIII + 8; anal rays 6-7 (usu- of dragonets with some researchers following the ally 7); all dorsal rays branched; anal rays scheme of Nakabo (1982) in which 19 genera are unbranched (except last branched at base); preop- recognized, while others subscribe to the classifica- ercular spine with very small main tip and 1-3 tion of Fricke (1983), who recognized only nine recurved spinules along dorsal margin; colour in genera, placing the majority of species in Callion- life mainly bright red with black pectoral fin base, myus Linnaeus, 1758 and Synchiropus Gill, 1859. diffuse blackish zone on ventral half of side punc- The present paper describes a new species that we tuated by variable-sized white spots; male with first became aware of from photographs taken at moderately elevated spinous dorsal fin with four Bali, Indonesia, labelled as Synchiropus cf. mor- black oblique bands, alternating with narrower risoni, which appeared in Kuiter & Tonozuka bands of yellow and blue; female with shorter, (2001). Four specimens were eventually collected mainly black spinous dorsal fin. in September 2010 by the second author during Description: Dorsal rays VIII + 8, all soft rays deep (60-70 m) scuba dives at Cenderawasih Bay, unbranched; anal rays 7 (one paratype with 6), West Papua Province, Indonesia. The specimens unbranched except last ray branched at base; pec- include both sexes and clearly belong to a new toral rays 21; pelvic rays I,5; branched caudal rays taxon, closely related to S. morrisoni Schultz, 1960. 7 (one paratype with 6); principal caudal rays 9 (one paratype with 8); upper and lower procurrent MATERIALS AND METHODS caudal rays 3-4. The format of the new species description and Body elongate and slightly depressed, greatest methods of counting and measuring are the same depth 4.8 (4.6-4.8) in SL, and greatest width 1.0 as those utilised by Fricke (1981 and 1983). We (0.9-1.0) in greatest depth. Head slightly com- have adhered to Fricke’s generic classification pressed (male) or slightly depressed (female), rather than that of Nakabo (1982), primarily length 3.3 (2.8-3.2) in SL; snout shorter than orbit because of the former author’s in-depth treatment diameter, 3.7 (3.8-4.4) in head length; orbit diam- of Indo-Pacific species in the 1983 monograph and eter 2.5 (2.4-2.6) in head length; interorbital very subsequent publications. Counts and proportions narrow, 15.4 (18.0-22.0) in head length. Caudal appearing in parentheses apply to the paratypes if peduncle moderately long and slender, least depth different from the holotype. Proportional measure- 10.1 (10.6-11.8) in SL and length 6.7 (4.9-6.2) in ments of type specimens, expressed as percentage SL. Branchial opening sublateral in position. Pre- of the standard length, are provided in Table I. opercular spine length 3.6 (4.1-4.7) in head length; Type specimens are deposited at Museum Zoolog- preopercular spine with very small main tip and icum Bogoriense, Cibinong, Indonesia (MZB) and 1-3 recurved spinules along dorsal margin. the Western Australian Museum, Perth (WAM). Lateral line extending from preorbital region to about middle of third branched caudal fin ray (counted from dorsal margin) with short subor- Synchiropus tudorjonesi n. sp. bital and long preoperculo-mandibular branch, Red-back Dragonet two ventral branches before pectoral fin base, and Figs 1-2; Table I series of short dorsal branches along side of body; lateral lines of opposite sides of body intercon- Holotype: MZB 20019, male, 35.5 mm SL, nected by commissure across occipital region. oceanic patch reef, 02°25.830’S, 134°54.409’E, Snout to origin of first dorsal fin 2.9 (2.5-2.8), north of Tridacna Atoll, Cenderawasih Bay, West and origin of second dorsal fin 2.0 (1.8-2.0), both aqua vol. 18 no. 1 - 15 January 2012 10 Gerald R. Allen and Mark V. Erdmann in SL; first dorsal fin of male holotype moderately short, 1.3 (1.7-1.9) in head length. Caudal fin elevated, first spine 2.0, second spine 1.4, third slightly rounded, 3.1 (3.2-3.9) in SL. spine 1.3, and fourth spine 1.5, all in head length. Colour in life (Fig. 1): mainly bright mot- First dorsal fin of female paratypes lower, first spine tled red with black pectoral fin base, diffuse black- 2.2-3.7, second spine 2.1-3.1. third spine 1.9-2.5, ish zone on ventral half of side, both upper and and fourth spine 3.3-4.1, all in head length. First lower side punctuated by variable-sized white dorsal fin higher than second dorsal fin in male spots; lower side of head and breast whitish to yel- and female, first and last dorsal rays 1.7 (1.9-2.0) lowish (with small blue spots on male); first dorsal and 1.3 (2.1-2.4) respectively in head length, the fin of male with four black oblique bands, alter- last ray slightly elongated in male. Preanal length nating with narrower bands of yellow and blue; 1.7 (1.6) in SL; anal fin origin level with base of female with mainly black spinous dorsal fin; sec- third ray of second dorsal fin; length of last anal ray ond dorsal fin reddish with white and dark brown 1.3 (1.5-1.8) in head length. Prepelvic length 3.3 spots arranged in oblique bands; caudal fin with (3.0-3.3) in SL; pelvic fins relatively elongate and about 6-7 alternating brown and white to bluish extending to base of third anal ray when laid back, bars intermingled with white spots; anal fin black- 0.9 (0.9-1.0) in head length. Pectoral fins relatively ish; pelvic fins translucent with diffuse red and yel-

Fig. 1. Synchiropus tudorjonesi, underwater photograph of anesthetized freshly collected male holotype, 35.5 mm SL (upper fish), and female paratype, 25.2 mm SL, Cenderawasih Bay, Indonesia. Photo by G. R. Allen.

11 aqua vol. 18 no. 1 - 15 January 2012 A new species of Dragonet (Synchiropus: Callionymidae) from Indonesia lowish bands; pectoral fins translucent with large 2005; personal observations) with records from the reddish blotches on basal half. Japan (Izu and Ryukyu Islands), Micronesia (Palau Colour in alcohol (Fig. 2): overall yellow- to Marshall Islands), Fiji, Samoa, Chesterfield ish tan with six large dark brown blotches (bifur- Islands, Great Barrier Reef, Indonesia (Bali, cate ventrally) on lower half of side; first dorsal fin Sulawesi, and West Papua), Philippines (Luzon), of male with four dark brown oblique bands, alter- and Papua New Guinea (Madang). Both species nating with narrower bands of dark brown and exhibit similar meristic and morphometric fea- white; female with mainly black spinous dorsal fin; tures. However, the adult male of S. morrisoni has soft dorsal fin whitish with 3-4 dark brown streaks a much taller first dorsal fin, which may reach on each fin membrane; caudal fin whitish with 2- twice the height of the second dorsal fin (or to 2.5 3 (only darker bars seen in life remain evident) hor- in SL) compared to the much lower fin (4.2 in SL) izontally elongate streaks on each fin membrane, of S. tudorjonesi. There is also a pronounced differ- most prominent on basal half; anal fin dark to ence in the colour pattern of the male’s first dorsal dusky brown except narrowly whitish at base; fin: that of S. morrisoni has numerous (generally pelvic and pectoral fins whitish with dark brown more than eight), mainly vertically oriented dark patch on upper half, extending dorsally on “shoul- brownish bands compared to the four black bands der” where it encircles pale spot just above fin. of S. tudorjonesi. Although the general colour pat- Remarks: The new species appears to be most tern of the body is similar for the two species, that closely related to Synchiropus morrisoni (Fig. 3), of S. morrisoni is generally more brownish than red, which ranges widely in the western Pacific (Randall and tends to be more strongly mottled. Finally

Fig. 2. Synchiropus tudorjonesi, preserved male holotype, 35.5 mm SL (upper fish), and female paratype, 25.2 mm SL, Cenderawasih Bay, Indonesia. Photo by G. R. Allen. aqua vol. 18 no. 1 - 15 January 2012 12 Gerald R. Allen and Mark V. Erdmann

Table I. Proportional measurements of type specimens of Synchiropus tudorjonesi as percentage of the standard length.

Character Holotype Paratype Paratype Paratype MZB WAM WAM WAM 20019 P.33368 P.33357 20020

Sex male female female female Standard length 35.5 25.2 24.4 21.2 Body depth 20.8 21.0 21.7 20.8 Body width 21.7 22.6 23.4 21.2 Head length 30.4 35.7 33.6 31.1 Snout length 8.2 9.5 7.8 7.1 Eye diameter 12.4 13.9 13.9 12.3 Interorbital width 2.0 2.0 1.6 1.4 Preopercular spine length 8.5 7.5 8.2 6.6 Caudal peduncle depth 9.9 8.7 9.4 8.5 Caudal peduncle length 14.9 20.2 16.4 16.0 Urogenital papilla length 0.8 0.8 0.0 0.0 Predorsal (1) length 34.6 39.7 39.8 35.4 Predorsal (2) length 51.0 55.6 53.3 50.0 Preanal length 58.0 62.3 62.7 61.3 Prepelvic length 30.7 33.3 32.0 30.7 First dorsal spine length 15.5 12.3 15.2 8.5 Second dorsal spine length 22.3 11.9 16.4 9.9 Third dorsal spine length 23.9 14.7 17.6 12.3 Fourth dorsal spine length 19.7 8.7 10.2 9.4 First dorsal ray length 17.5 18.3 17.6 15.6 Last dorsal ray length 24.2 17.9 16.0 12.7 Last anal ray length 23.4 19.8 18.4 20.3 Pectoral fin length 23.1 19.0 17.6 18.4 Pelvic fin length 33.0 35.3 36.1 34.4 Caudal fin length 32.7 31.7 29.9 25.9

Fig. 3. Synchiropus morrisoni, male, 57.0 mm SL, Okinawa, Ryukyu Islands. Photo by J. E. Randall.

13 aqua vol. 18 no. 1 - 15 January 2012 A new species of Dragonet (Synchiropus: Callionymidae) from Indonesia there is also a difference in habitat with S. mor- vey, which was completed under Teluk Cen- risoni usually occurring at depths between about drawasih National Park license No. SI.1029/IV- 12-33 m (Randall 2005; personal observations) 14/TEK/2010. We sincerely thank the Siebel Fam- compared to 50-70 m for the new species. ily Charitable Foundation, Rodney and Nancy Habitat and distribution: The new species was Chiamulon, and Dan Cohen for their generous collected from 70 m depth on reefs of Cender- financial support of this survey, and also the awasih Bay that are situated well offshore in clear Indonesian Institute of Sciences (specifically Dr. water, oceanic conditions. The type specimens Suharsono and Ibu Rianta Afriadi) for loaning the were each collected in a similar microhabitat that specimens for study. Finally, we thank the captain consisted of scattered coral and sponge outcrops and crew of the MV TemuKira for their tireless with interspersed rubble substrate; when startled, efforts in supporting an exhaustive dive schedule. the animals would invariably seek shelter under the nearest outcrop. Individuals from Tulamben, Bali, Indonesia were also reported (as Synchiropus cf. REFERENCES morrisoni) by Kuiter and Tonozuka (2001) from a FRICKE, R. 1981. Revision of the genus Synchiropus depth of 50 m. (Teleostei: Callionymidae). Theses Zoologicae 1: 1-194. Etymology: The new species is named tudorjonesi FRICKE, R. 1983. Revision of the Indo-Pacific genera and species of the dragonet family Callionymidae (Teleostei). after Paul Tudor Jones, in honour of his dedication Theses Zoologicae 3: 1-774. and selfless service to the United States National GILL, T. N. 1859. On the genus Callionymus of authors. Fish and Wildlife Foundation (NFWF). Under his Proceedings of the Academy of Natural Sciences of Philadel- strong leadership as chairman from 2006 to 2009, phia 11: 128-130. the NFWF has grown into a highly respected con- KUITER, R. H. & TONOZUKA, T. 2001. Indonesian Reef servation institution and expanded its programs Fishes. Zoonetics, Seaford, Victoria, Australia, 893 pp. from a national focus to a global one, including pro- LINNAEUS, C. 1758. Systema Naturae, Ed. X. (Systema nat- tection of the Bird’s Head Seascape reefs which, this urae per regna tria naturae, secundum classes, ordines, genera, species, cum characteribus, differentiis, syn- beautiful dragonet species calls home. Through onymis, locis. Tomus I. Editio decima, reformata.) these efforts, Mr. Tudor Jones has forever left his Holmiae, 824 pp. mark on the noble pursuit of nature preservation. NAKABO, T. 1982. Revision of genera of the dragonets (Pisces: Callionymidae). Publications of the Seto Marine ACKNOWLEDGEMENTS Biological Laboratory 27 (1/3): 77-131. We thank the Balai Besar Taman Nasional Teluk RANDALL, J. E. 2005. Reef and Shore Fishes of the South Cenderawasih (particularly Christina Matakupan Pacific. University of Hawai’i Press, Honolulu, 707 pp. SCHULTZ, L. P. 1960. Family Callionymidae. In: Fishes of and Titus Wemiyaupea) for their support and par- the Marshall and Marianas islands. Volume 2. Families ticipation in the field survey that uncovered this from Mullidae through Stromateidae. (Eds. L. P. Schultz, species. We also thank Hamid Toha and the State W. M. Chapman, E. A. Lachner & L. P. Woods). Bulletin University of Papua for their assistance in the sur- of the United States National Museum 202: 1-438.

aqua vol. 18 no. 1 - 15 January 2012 14 aqua, International Journal of Ichthyology

Reproductive Biology of the Silky Shark Carcharhinus falciformis (Chondrichthyes: Carcharhinidae) off the west coast of Baja California Sur, Mexico

E. Mauricio Hoyos-Padilla1, B. Patricia Ceballos-Vázquez2 and Felipe Galván-Magaña2*

(1) Pelagios-Kakunja, A. C. Sinaloa 1540, La Paz, Baja California Sur, Mexico. 2) Centro Interdisciplinario de Ciencias Marinas (CICIMAR-IPN). Av. Instituto Politécnico Nacional s/n Col. Playa Palo de Santa Rita, P.O. Box 592, La Paz, Baja California Sur, Mexico. C. P. 23096. E-mail: [email protected]; [email protected]; *Corresponding author: [email protected]

Received: 14 January 2011 – Accepted: 12 August 2011

Abstract Geschlechterverhältnis zwischen Weibchen und Männchen The silky shark Carcharhinus falciformis population has betrug 1:0,6. Nimmt man die Zunahme des Durchmessers declined drastically in the last few years, due to extensive der Eier in den Eierstöcken und der Eileiterdrüsen sowie das by-catch in tuna purse-seine and longline fisheries in the Vorhandensein von Eiern im Uterus oder die Entwicklung eastern Pacific Ocean. No information exists on the repro- von Embryos zum Maßstab, muss man zu dem Schluss ductive biology of this species in the eastern Pacific Ocean kommen, dass die Geschlechtsreife der Weibchen bei einer to support fishery or conservation management. A total of Länge von 180 cm TL eintritt; Männchen werden bei 182 295 silky sharks were analyzed, with 179 females captured, cm TL geschlechtsreif, gemessen an der Entwicklung des ranging between 88 and 316 cm TL and 116 males, rang- Klammerorgans und der Anwesenheit von Samenklumpen. ing between 142 and 260 cm TL. The sex ratio of females Die kurzzeitige Speicherung von Samen, wie man sie bei to males was 1: 0.6. The increase in of oocytes instead of Weibchen feststellen konnte, dürfte bei Arten, bei denen die Geschlechter getrennt und im freien Wasser leben, von ovarian egg and oviducal gland diameters as well as the Vorteil sein, sie steigert den Fortpflanzungserfolg. Bei den presence of uterine eggs or developing embryos indicated 20 untersuchten trächtigen Weibchen fanden sich zwei bis that female maturation occurred at about 180 cm TL; neun, durchschnittlich fünf Embryos. Weibchen mit 6-8 while clasper development and the presence of sperm cm TL großen Embryos konnten im Juli und September clumps indicated that males matured at about 182 cm TL. festgestellt werden, während 20-30 cm TL lange Embryos The short-term sperm storage found in females could be zwischen September und November vorkamen. Ein an advantage for species that present sexual segregation Weibchen mit voll entwickelten Embryos (80 cm) konnte and live in open waters, increasing reproductive efficiency. Ende Juni gefangen werden, was auf eine Trächtigkeitsdauer In the 20 gravid females examined, the average number of von 11-12 Monaten schließen lässt. embryos per female was five, with a range of 2-9 embryos. Females with embryos 6-8 cm TL were observed in July Résumé and September; whereas embryos 20-30 cm TL were La population du Carcharinus falciformis a drastiquement found from September to November. One female with full diminué ces dernières années à cause de la capture involon- term embryos (80 cm) was captured at the end of June sug- taire en sennes à thons et de la pêche à longue ligne dans gesting an 11-12 month gestation period. le Pacifique est. On ne dispose pas d’informations sur la biologie reproductive de cette espèce dans le Pacifique est Zusammenfassung pour assister la pêche oula préservation. Un total de 295 Die Bestände des Glatthais Carcharhinus falciformis sind in individus ont été examinés : 179 femelles, de 88 à 316 cm den letzten Jahren drastisch zurückgegangen, verursacht de LT, et 116 mâles, de 142 à 260 cm de LT. Le sex ratio durch die Tunfischerei mit Ringwadennetzen und Lang - des femelles par rapport aux mâles était de 1:0,6. L’aug- leinen im östlichen Pazifischen Ozean, bei der Glatthaie als mentation du diamètre de l’œuf ovarien et de la glande de Beifang in die Netze geraten. Über die Fortpflanzungsbiolo- l’oviducte, ainsi que la présence d’œufs utérins ou d’em- gie dieser Art im östlichen Pazifik gibt es bisher keine Infor- bryons en développement ont indiqué que la maturité de mationen, mit denen Fischereimanagement oder Schutz- la femelle se produisait à environ 180 cm de LT ; d’autre maßnahmen unterstützt werden könnten. Insgesamt wur- part, le développement du ptérygopode et la présence de den 295 Glatthaie analysiert; 179 der gefangenen Tiere paquets de sperme montrent que les mâles sont mûrs à près waren Weibchen zwischen 88 und 316 cm Gesamtlänge de 182 cm de LT. Le stockage de courte durée de sperme (TL), 116 Männchen zwischen 142 und 260 cm TL. Das trouvé chez des femelles pourrait être un avantage pour des

15 aqua vol. 18 no. 1 - 15 January 2012 Reproductive Biology of the Silky Shark Carcharhinus falciformis (Chondrichthyes: Carcharhinidae) off the west coast of Baja California Sur, Mexico espèces qui pratiquent une ségrégation sexuelle et vivent en line fisheries, but is also taken in gill nets off the eau libre, en augmentant l’efficacité reproductive. Pour les west coast of Mexico, representing one of the ten 20 femelles pleines examinées, le nombre moyen d’em- most important shark species in captures from bryons par femelle était de cinq, dans une fourchette de 2-9 Mexican Pacific waters (Castillo-Geniz et al. embryons. Des femelles avec de embryons de 20-30 cm de LT on été trouvées de septembre à novembre. Une femelles 2010). Despite its high abundance abundance in porteuse d’embryons arrivés à terme (80 cm) a été capturée the region, there are few data concerning C. falci- fin juin, ce qui suppose une gestation de 11-12 mois. formis’ biology and studies focused on its repro- ductive biology are necessary to design and imple- Sommario ment strategies for the regulation of its fishery. Negli ultimi anni le popolazioni di squalo seta Carcharhi - Some published information exists on reproduc- nus falciformis del Pacifico orientale si sono ridotte drastica- tion of this species, limited to the studies of Gilbert mente a causa delle numerose catture accidentali con reti a & Schlernitzauer (1965, 1966), Branstetter (1987) circuizione e con palamito. Non esistono dati sulla biologia and Bonfil et al. (1993). The only study done on riproduttiva di questo squalo nell’oceano Pacifico orientale tali da sostenere politiche di controllo della pesca o di con- C. falciformis in Mexican waters is that of Bonfil et servazione della specie. Complessivamente sono stati analiz- al. (1993); however, the information is from the zati 295 esemplari di squalo seta, 179 femmine di ta glia Mexican Atlantic Ocean. Silky shark biology from compresa tra gli 88 e i 316 cm TL e 116 maschi di ta glia this area has been briefly discussed in a bachelor’s compresa tra i 142 e i 260 cm TL. Il rapporto femmi ne: thesis (Cadena 2001). The aim of the present study maschi era di 1: 0.6. L’incremento del diametro delle uova e is to determine the reproductive parameters and delle ghiandole dell’ovidotto e la presenza di uova ute rine o morphology of the reproductive tract and sperm embrioni in fase di sviluppo indicano che la fem mi na rag- storage of both sexes in C. falciformis from the east- giunge la maturazione a circa 180 cm TL; lo sviluppo dello pterigopodio e la presenza di masse di sperma in dicano che ern Pacific Ocean. These data would allow the il maschio matura a circa 182 cm TL. Il man tenimento stock assessment of species harvested by fisheries dello sperma solo per brevi periodi nella fem mina potrebbe and the ecological risk assessment of species caught essere un vantaggio per la specie che mo stra segregazione dei as bycatch (Walker 2004). sessi e abita acque aperte, incrementando l’efficienza ripro- duttiva. Nelle 20 femmine gravide esaminate il numero MATERIALS AND METHODS medio di embrioni per femmina era pari a cinque, con un From 2000 to 2002, silky sharks were obtained intervallo di 2-9 embrioni. Femmine con embrioni di 6-8 from commercial catches off Punta Lobos and cm TL erano osservate in luglio e settembre, mentre embrio- ni di 20-30 cm TL erano rinvenuti da settembre a novem- Punta Belcher (Fig. 1), the two major shark fishing bre. Una femmina con embrioni allo stadio finale (80 cm) è camps in Baja California Sur, Mexico. Specimens stata catturata alla fine di giugno suggerendo un periodo di from Punta Lobos were caught by longlines while gestazione di 11-12 me si. in Punta Belcher they were caught with gill nets. Total length was measured (Compagno 1984). INTRODUCTION Clasper length was measured from the insertion of The silky shark, Carcharhinus falciformis, is a cos- the inner corner of the pelvic fin to its distal tip. mopolitan species found throughout tropical and The meager information available for shark species subtropical waters. In the eastern Pacific it ranges and critical examination of recently acquired spec- from southern Baja California to Peru. It is an imens demonstrates that, even though there are abundant offshore oceanic and epipelagic tropical shared reproductive characters throughout the shark. It occasionally occurs inshore where the orders, there are also differences in the reproduc- water is as shallow as 18 m; in the open ocean it tive biology between the various genera and species occurs from the surface down to at least 500 m (Castro 1996). Males were considered mature depth. Water temperatures of 23° to 24° C have when claspers were completely calcified and the been recorded where it occurs (Compagno 1984). riphiodon could be spread open (Springer 1960; In the eastern Pacific Ocean the silky shark is cap- Clark & von Schmidt 1965). Females were consid- tured by longline and occasionally with gillnets ered mature when they had at least one of the fol- (Au 1991), and makes up to 30% of all captured lowing characteristics: gravidity, ripe ovarian eggs, sharks; however, this species has declined drasti- oviducal gland width over 20 mm in diameter, or a cally in recent years, due to by-catch from tuna well-developed uterus. Most of the time checking purse-seiners in the eastern Pacific Ocean (Watson for uterine scars in the uterus was undertaken for 2008). This species is common in the pelagic long- most of the females, to avoid taking into account aqua vol. 18 no. 1 - 15 January 2012 16 E. Mauricio Hoyos-Padilla, B. Patricia Ceballos-Vázquez and Felipe Galván-Magaña aborted females; sometimes fishermen allowed us ratio was 1F:0.6M. The range in total length for to take samples just while they are preparing the females was 88-316 cm (average 180 cm TL), and sharks for transportation, so in a few cases it was for males the range was 142-260 cm TL (average impossible to be sure of the presence of these scars. 182 cm TL). Specimens 176 to 225 cm TL com- Additionally, maturity was corroborated in males prised 76% of the total captures. by the presence of sperm clumps in the seminal Morphological and histological analysis: males vesicle (Pratt & Tanaka 1994); whereas in females The testes in the silky shark are almost totally sur- it was corroborated by the presence of sperm in the rounded by the epigonal organ except in the distal oviducal gland. The reproductive system was region. They are paired, elongated and dorsoven- removed and fixed in buffered 10% formalin. Sec- trally flattened organs (Fig. 2), which are attached tions 5-6 µm thick were prepared using the con- to the dorsal wall of the abdominal cavity by a ventional histological process with haematoxylin- mesorchium. Depending on the size of the adult, eosin stain (Castro & Wourms 1993) to analyze the testes range from 17 to 33.5 cm in length. the gonads under the microscope. Seminiferous follicle development spreads from a germinal zone across the diameter of the testicles to RESULTS the opposite wall, where the efferent ductules form Seasonal abundance, sex ratio and size a manifold collection system to receive spermato- Silky sharks are seasonal in commercial catches of zoa (Fig. 3). The efferent ductules continue in a the west coast of Baja California Sur from June to series of fine tubules and communicate with the November (summer and autumn), with a maxi- head of the epididymis. mum frequency in September. A total of 179 The epididymides are convoluted, unpigmented females and 116 males were examined. The sex and digitiform (Fig. 2). In adults, each epididymis

Fig. 1. Location of the fishing camps Punta Belcher and Punta Lobos in the west coast of Baja California Sur, Mexico.

17 aqua vol. 18 no. 1 - 15 January 2012 Reproductive Biology of the Silky Shark Carcharhinus falciformis (Chondrichthyes: Carcharhinidae) off the west coast of Baja California Sur, Mexico is approximately 25 cm long and 2 cm wide. The of the Leydig gland has two kinds of cells: ciliated tubules of this organ are divided in two types: Ley- and secretory (Fig. 4a). The epididymis tubule has dig glands and epididymis tubules. The epithelium a pseudostratified columnar epithelium with cilia and individual spermatozoa scattered in the lumen (Fig. 4b). In addition to sperm, three types of par- ticulate matter occur in the lumen of the ducts: Sertoli cell bodies, Sertoli cytoplasts, and Leydig gland bodies. The role of these components in sperm aggregation or maintenance is unknown (Hamlett 1999). Towards the terminal end of the epididymis the ductus deferens are formed (Fig. 2). In C. falciformis, organized masses of naked sperm bound with a cohesive matrix forming ovoid to sub-spherical masses were found in the seminal vesicle (Fig. 2). These ovoid sperm aggregates (96 to 310 µm) were found as a single layer of radially aligned sperm clumps organized around a core of eosinophilic material (Fig. 5) known as single layer spermatozeugmata (Pratt & Tanaka 1994). Morphological and histological analysis: females In C. falciformis, only the right ovary is func- tional, as in most carcharhinids (Pratt 1988). It is located at the distal surface of the epigonal organ, at the forward end of the abdominal cavity (Fig. 6), and attaches to the dorsal wall of the abdominal cavity by a mesovarium. The ovary in adults ranged from 9 to 25 cm long and from 2.6 cm to 9.6 cm wide. Ovulating females carried 8-11 oocytes 30- 35 mm in diameter. The silky shark ovary con- Fig. 2. Male reproductive system. Epididymis (E), testis tained corpora atretica in various stages of develop- (T), deferens ducts (DD), epigonal organ (EO) and sperm ment. In 86 ovaries, we found these structures to sacs (SS). range from 4-34 mm in diameter. Post-ovulatory follicles (corpora lutea) from 6 to 33 mm were found in the ovaries of pregnant females. The rela- tive development of the corpora lutea and corpora atretica and their function remains unknown (Dodd 1983). The paired oviducts are slender seg- ments, which emerge from the peritoneal cavity by an ostium in the falciform ligament (Fig. 6). The upper portion of each oviduct is narrow and short, and leads into the oviducal gland. This gland is heart-shaped, from 13 to 50 mm in diameter. Externally, each gland resembles a symmetrical white organ with two horns on the lateral anterior surface (Fig. 6). Sperm storage was found in only one of 50 glands of analyzed mature females. The (GZ) spermatozoa were located in the thin-walled ← tubules of the oviducal gland around the curves of the lumen. The tubules are formed by two differ- ent types of cells, secretory and ciliated. The sperm Fig. 3. Cross section of the testis showing the diametric was loosely packed in tubules with well-stained development of follicles. Germinal zone (GZ). heads up current (Fig.7). aqua vol. 18 no. 1 - 15 January 2012 18 E. Mauricio Hoyos-Padilla, B. Patricia Ceballos-Vázquez and Felipe Galván-Magaña

a b

Figure 4a-b. Micrographs of Leydig gland (a) and epididymis tubule (b).

The uterus is divided in two parts, one of which Size at maturity is designated as the third membrane store chamber. Males: Development of the testis is gradual; at In the oviducal gland, the fertilized egg is enclosed about 180 cm TL it can reach a length of 25 cm in a membrane; it then passes through the third (Fig. 8). Sharks with uncalcified claspers had testes membrane store chamber and enters the uterus. in early stages of spermatogenesis and a small The third membrane remains around the embryo amount of sperm present in the lower epididymis. until it reaches birth size. The uterus size increases The claspers begin to elongate at about 160 cm TL with the size of the developing embryo. In females and become calcified at 179 cm TL. All specimens with uterine eggs, the uterus is 20-22 cm long; in larger than 181 cm TL were mature, with sperm females with small embryos (6-8 cm TL), the aggregates present in the seminal vesicle (Fig. 9). uterus is 33-34 cm long; and in term females (80 The size at maturity L50 (with 95% CI) in males cm embryos), the uterus reaches almost 50 cm long occurred at 182 (180-182) cm TL. and a width of about 15 cm. Females: The size of the oviducal gland started to increase when females reached 180 cm TL (Fig. 10). In juveniles under 170 cm TL the gland mea- sured 5-10 mm in width; whereas in females 180 cm TL it measured 20-40 mm in width. Immature females had undeveloped oocytes 6-10 mm in maximum diameter. The oocytes did not start growing until the females reached 180 cm TL (Fig. 11). The size at maturity L50 (with 95% CI) in females occurred at 180 (179-180) cm TL. Larger females were also seen to be carrying ripe oocytes 20-43 mm in diameter. The presence of ovulatory females with mating bites on their bodies or with sperm in their uteri was very sporadic. Only two recently inseminated females were caught during our study, on 31 August 2001 and 31 July 2002. Fig. 5. Micrographs showing the sperm clump in males. One of them had oocytes 32 mm in diameter.

19 aqua vol. 18 no. 1 - 15 January 2012 Reproductive Biology of the Silky Shark Carcharhinus falciformis (Chondrichthyes: Carcharhinidae) off the west coast of Baja California Sur, Mexico

Another female caught on 24 November 2000 bore DISCUSSION mating scars and oocytes 35 mm in diameter. In Carcharhinus falciformis begins its northward our sample of 20 gravid females, the average num- migration in June. The schools stay off the west ber of embryos per female was five, with a range of coast of Baja California Sur from June to October, 2-9 embryos. Ovulating females with uterine eggs arriving off Punta Belcher and Punta Lobos in and embryos 6-8 cm TL were observed in July and August, although most of the adults arrive in Sep- September, whereas embryos 20-30 cm TL were tember and October (Fig. 12). The timing of the C. found from September to November. Only one falciformis captures was consistent with changes in female with full term embryos (80 cm) was cap- the surface water temperature. Gómez & Vélez tured at the end of June. (1982) recorded that the water temperature of the California current increases to 20°C to 25°C in summer. Carcharhinus falciformis leave the west coast of Baja California Sur in early fall and migrate southward when the sea surface water temperature decreases below 20°C. It is known that the increase in water temperature influences the migrations of other carcharhinids such as C. isodon and C. lim- batus off the east coast of the U.S.A. (Castro 1993, 1996). Most of the C. falciformis examined along the west coast of Baja California Sur were large speci- mens (176-225 cm TL). Cadena (2001) found sim- ilar sized sharks (170-220 cm TL) in the Gulf of California. Juveniles of C. falciformis have been recorded in coastal waters of south-west Mexico. Chong-Robles (2003) found gravid females carrying term embryos and neonates (50-83 cm TL) in March and May off the Oaxaca, Mexico, coast; whereas Ronquillo (1999) found gravid females in Chiapas, Mexico, during June. It appears that C. fal- ciformis use the coastal waters of south-west Mexico as nursery grounds and migrate to the north in the summer to feed and possibly to copulate. Villatorio & Rivera (1994) reported a 1F:1M sex ratio. Del Fig. 6. Female reproductive system. Ovary (O), oviducal Rosario (1998) and Ronquillo (1999) reported the gland (OG), uterus (U), and epigonal organ (EO). same ratio for Guatemala and Chiapas respectively. Bonfil et al. (1993) suggested a 1F:1M ratio for C. falciformis in nature. Their results differ from our findings (1F:0.6M), although those studies corre- spond to annual samplings and the presence of C. falciformis off the western coast of Baja California Sur is seasonal, occurring from July to October. Maturation in males is reached at about 182 cm TL and it involves the gradual development of the testes, the calcification of the claspers and the pres- ence of sperm aggregates in the seminal vesicle. The testes development (diametric) and spermato- genesis in C. falciformis are similar to those found in the catshark Scyliorhinus canicula (Dodd 1983). According to Hamlett (1999), in all elasmobranchs thus far examined the fundamental details of sem- Fig. 7. Micrographs showing the sperm storage in the iniferous follicle development and spermatogenesis oviducal gland. are similar. The sperm clumps found in C. falci- aqua vol. 18 no. 1 - 15 January 2012 20 E. Mauricio Hoyos-Padilla, B. Patricia Ceballos-Vázquez and Felipe Galván-Magaña

Fig. 8. Total length versus testes length.

Fig. 9. Total length versus clasper length and degree of calcification. The cross corresponds to the males with fully calcified claspers and spermatozeugmata in the seminal vesicle.

21 aqua vol. 18 no. 1 - 15 January 2012 Reproductive Biology of the Silky Shark Carcharhinus falciformis (Chondrichthyes: Carcharhinidae) off the west coast of Baja California Sur, Mexico

Fig. 10. Total length versus oviducal gland length.

Fig. 11. Total length versus egg diameter. aqua vol. 18 no. 1 - 15 January 2012 22 E. Mauricio Hoyos-Padilla, B. Patricia Ceballos-Vázquez and Felipe Galván-Magaña

Table I. Size at maturity of silky shark Carcharhinus falci- the Tasman Sea. A possible explanation in relation formis in different parts of the world. to differences in the maturity size is that individu- als from the eastern Pacific Ocean are smaller than Author and year Region Size at maturity those from the Indian Ocean. This has been TL (cm) recorded for other carcharhinids like the black tip Strasburg (1958) Central Pacific ♀213-216 shark, C. limbatus. Castro (1996) stated that C. INDIAN OCEAN limbatus in the western North Atlantic differs sig- Fourmanoir (1961) Madagascar ♀248-269 ♂240 ♀ ♂ nificantly in numerous traits such as size, mark- Stevens (1984) Aldabra Atoll 216 239 ings, age at maturity, brood size and other charac- SOUTH PACIFIC Stevens (1984) Tasman Sea ♀202-208 ♂214 ters from the Pacific specimens. EAST PACIFIC We observed the absence of a defined seasonality Bonfil et al. (1993) Campeche Bank ♀232-246 ♂225 for C. falciformis reproduction off the west coast of Del Rosario (1998) Pacific coast of ♀176 ♂178 Baja California Sur. The sperm found inside one Guatemala oviducal gland belonged to a pregnant female with Ronquillo (1999) Gulf of Tehuantepec, ♀180 intrauterine eggs. The arrangement of the sperm Mexico inside the tubules coincides with the “short-term Cadena (2001) Gulf of California, ♀180 ♂180 Mexico sperm storage” classification defined by Pratt Present study West Coast of ♀180 ♂182 (1993) and is found in sharks in which ovulation is Baja California Sur, prolonged over several weeks. Sperm storage is an Mexico advantage for species that presents sexual segrega- tion and lives in open waters, increasing the effi- formis (single layered spermatozeugmata), have ciency in reproduction. Strasburg (1958) stated been found in other carcharhinids like the black tip that the brood size of C. falciformis was two to shark Carcharhinus limbatus and the sandbar shark eleven embryos. Gilbert & Schlenitzauer (1966) Carcharhinus plumbeus (Pratt & Tanaka 1994). We report one female with nine embryos in the confirmed that the presence of sperm aggregates is Atlantic Ocean. Cadena (2001) gave a range of 1- a more reliable indicator of maturity than clasper 12 young for specimens seen off the Gulf of Cali- condition because the presence of sperm aggregates fornia. In the 20 gravid females recorded in our ensures that if the shark copulates there is a better study, the average number of embryos per female probability of fecundation. We acknowledge that was five, with a range of 2-9 embryos. Unfortu- the use of the clasper calcification criterion instead nately characteristics such as low fecundity and a of the presence of sperm aggregates is more practi- late age of sexual maturation, leave species like C. cal and economical (but the sperm aggregation falciformis vulnerable to overexploitation. method is more accurate by far). Maturation in females is reached at about 180 cm TL and is marked by the growth of the oviducal gland, enlargement of the uteri and the ripening of the oocytes. This size at maturity corresponds with the sizes recorded for C. falciformis in the Mexican Pacific and Guatemala (Del Rosario 1998; Ron- quillo 1999; Cadena 2001). Although C. falci- formis is considered a circumtropical species (Cas- tro 1983), the size at maturity is different in differ- ent areas (Table I). Fourmanoir (1961) found for Madagascar that males mature at about 240 cm TL and females at about 248-269 cm TL. At Aldabra Atoll, Indian Ocean, Stevens (1984) found that males mature at about 239 cm TL and females at about 216 cm TL. In the Pacific Ocean, Strasburg (1958) recorded 12 pregnant females of 213-236 cm TL and Stevens (1984) recorded mature males Fig. 12. Silky shark, caught by artisanal fishermen, Punta at 214 cm TL and females at 202-208 cm TL for Lobos, Baja California Sur, México. Photo by M. Carrera.

23 aqua vol. 18 no. 1 - 15 January 2012 Reproductive Biology of the Silky Shark Carcharhinus falciformis (Chondrichthyes: Carcharhinidae) off the west coast of Baja California Sur, Mexico

ACKNOWLEDGEMENTS de la biología y pesquería del tiburón blanco Carcharhi- We would like to thank the Instituto Politécnico nus falciformis (Bibron, 1839) de las costas del Pacífico de Na cional (SIP) for funding the project “Trophic Guatemala. Bachelor´s Thesis. Universidad de San Carlos eco lo gy of sharks and mantas in the lower Gulf of de Guatemala, CEMA, Guatemala. 74 p. DODD, J. M. 1983. Reproduction in cartilaginous fishes California”; the fishermen of Punta Belcher, Punta (Chon drichtyes). Pp. 31-86. In: Hoar, W. S. & D. J. Lobos and Las Barrancas for their help during sam- Ran dall (Eds) Fish Physiology. Academic Press, Inc. New pling; and the Fish Ecology and Invertebrates Lab- York, E.U.A. oratories from CICIMAR. B.P.C.V. and F.G.M. FOURMANOIR, P. 1961. Requins de la cote ouest de Mada- received grants from SIBE (COFAA), EDI (IPN), gascar. Memories Institute of Science Madagascar 4: 1-81. and SNI-CONACYT. GILBERT, P. W. & SCHLERNITZAUER, D. A. 1965. 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Uni- WALKER, T. I. 2004. Chapter 13. Management measures. versidad del Mar, Puerto Ángel, Oaxaca. 66 p. In: Technical Manual for the Management of Elasmo- CLARK, E. & VON SCHMIDT, K. 1965. Sharks of central gulf branchs (Eds. Musick, J. A. & Bonfil R.): 285-32. Asia coast of Florida. Bulletin of Marine Science 15: 13-83. Pacific Economic Cooperation Secretariat, Singapore. COMPAGNO, L. J. V. 1984. Sharks of the world. An anno- WATSON, J. T., ESSINGTON, T. E., LENNERT-CODY, C. E. & tated and illustrated catalogue of shark species known to HALL, M. A. 2008. Trade-offs in the design of fishery clo- date. Part 2. Carchariniformes. FAO Fisheries Synopsis sures: Management of silky shark bycatch in the Eastern 125 (4): 251-663. Pacif ic Ocean tuna fishery. Conservation Biology 23 (3): DEL ROSARIO, R. C. 1998. Contribución al conocimiento 626-635. aqua vol. 18 no. 1 - 15 January 2012 24 aqua, International Journal of Ichthyology

A new species of Odontanthias Bleeker (Perciformes: Serranidae: Anthiinae) from Mona Passage off Puerto Rico, the first record of the genus from the Atlantic Ocean

William D. Anderson, Jr.1 and Graciela García-Moliner2, 3

1) Grice Marine Biological Laboratory, College of Charleston, 205 Fort Johnson, Charleston, South Carolina 29412-9110, USA. E-mail: [email protected] 2) Caribbean Fishery Management Council, 268 Muñoz Rivera Avenue, Suite 1108, San Juan, Puerto Rico, 00918-1920, USA. E-mail: [email protected] 3) Department of Marine Sciences, University of Puerto Rico, Mayagüez, PO Box 9000, Mayagüez, Puerto Rico 00681, USA.

Received: 21 April 2011 – Accepted: 27 August 2011

Abstract Odontanthias para la región Indo-Pacífica, pero éste es el Odontanthias hensleyi, a new species of anthiine serranid primer reporte de la presencia de una especie de dicho fish, is described from four specimens collected off the Género en el Océano Atlántico. La nueva especie se dis- west coast of Puerto Rico in Mona Passage. This is the first tingue por las siguientes características: quince rayos blan- Atlantic record for the genus Odontanthias, heretofore dos en la aleta dorsal, dieciocho rayos en las aletas pec- known from 13 Indo-Pacific species. The new species is torales, el parcho del diente vomerino de forma subcuad- distinguished from all other species of Odontanthias by the rangular o de diamante y sin una prolongación posterior, following combination of characters: 15 soft rays in the los lóbulos de las aletas pélvica y caudal pronunciados y dorsal fin, 18 pectoral fin rays, vomerine tooth patch sub- una coloración particular (franjas brillantes amarillas a quadrangular to diamond-shaped without posterior pro- ambos lados de la cabeza y las aletas pélvica, anal y caudal longation, pelvic fin and caudal fin lobes very well pro- del mismo amarillo). duced, and coloration (side of head with two bright yellow stripes; pelvic, anal and caudal fins bright yellow). Résumé Odontanthias hensleyi, une nouvelle espèce de Serranidé, Zusammenfassung est décrit sur base de quatre spécimens collectés au large de Auf der Grundlage von vier Exemplaren, die man vor der la côte ouest de Porto Rico, dans le canal de la Mona. C’est Westküste von Puerto Rico in der Mona-Passage gefangen la première mention atlantique du genre Odontanthias, hat, wird Odontanthias hensleyi als neue Art der Fahnen- connu jusqu’ici par 13 espèces de l’Indo-Pacifique. La nou- barsche (Serranidae: Anthiinae) beschrieben. Es handelt velle espèce se distingue de toutes les autres espèces sich um den ersten Nachweis der Gattung aus dem d’Odontanthias par la combinaison des caractéristiques Atlantik, von der man bisher 13 indo-pazifische Arten suivantes : 15 rayons mous dans la dorsale, 18 rayons dans kennt. Die neue Art lässt sich von allen anderen Odontan- la pectorale, une zone de dents du vomer subquadrangu- thias-Arten durch die Kombination folgender Merkmale laire à diamantiforme, sans prolongement postérieur, des unterscheiden: 15 Weichstrahlen in der Rückenflosse, 18 lobes de pelvienne et de caudale très allongés et la col- Brustflossenstrahlen, Vomerzahnabschnitt fast viereckig oration (le côté de la tête avec deux lignes jaune clair, pelvi- bis diamantförmig ohne Verlängerung nach hinten, sehr ennes, anale et caudale jaune clair). gut ausgebildete Bauch- und Schwanzflossen sowie durch die Farbgebung (zwei hellgelbe Streifen an den Kopfseiten; Sommario Bauch-, After- und Schwanzflossen hellgelb). Odontanthias hensleyi, è una nuova specie di castagnola descritta sulla base di quattro esemplari presi al largo della Resumen costa occidentale di Puerto Rico in Mona Passage. Si tratta Se describe una nueva especie de pez ánthido: Odontan- della prima specie del genere Odontanthias dell’Atlantico; thias hensleyi (Perciformes: Serranidae), a partir de cuatro infatti, questo genere era noto finora per 13 specie tutte especímenes recolectados en el Canal de La Mona, al oeste dell’Indo-Pacifico. La nuova specie si distingue da tutte le de Puerto Rico. Se conocen trece especies del Género altre del genere Odontanthias per la seguente combinazione

25 aqua vol. 18 no. 1 - 15 January 2012 A n. sp. of Odontanthias Bleeker from Mona Passage off Puerto Rico, the first record of the genus from the Atlantic Ocean di caratteri: 15 raggi molli nella dorsale, 18 raggi pettorali, Anderson & Baldwin (2000). Abbreviations denti vomerini in aree subquadrangolari o romboidali include: SL (standard length), UPRM (University senza prolungamenti posteriori, lobi delle pinne pelviche e of Puerto Rico, Mayagüez) and USNM (National della caudale molto allungati e colorazione (lati del capo Museum of Natural History, Smithsonian Institu- con due brillanti strie gialle; pinne pelviche, anale e cau- dale giallo acceso). tion, Washington, D.C.).

INTRODUCTION Odontanthias hensleyi n. sp. Randall & Heemstra (2006) reviewed the anthi- Euripos Jewelfish ine serranid genus Odontanthias Bleeker, 1873, (Fig. 1; Table I) considering it to include 13 Indo-Pacific species, two of which were new to science, and described a Holotype: USNM 400888, 155 mm SL, male; new anthiine genus, Meganthias. They (2006: 2) western North Atlantic, Mona Passage, off west noted that: “The generic classification of the fishes coast of Puerto Rico, 18°07’ N, 67°40’ W; 185 of the Anthiinae has been very confused, and this fathoms (338 meters); collected by Eugenio is certainly true for Odontanthias and related gen- Piñeiro Soler and Miguel A. Vargas, 20 July 2004. era.” Based on morphology the genera most closely Paratypes: UPRM 3793, 162 mm SL, & UPRM related to Odontanthias would appear to be Holan- 3794, 159 mm SL; Mona Passage, 18°24’ N, thias (with which it has been frequently syn- 67°40’ W; 188 fathoms (344 meters); 12 August onymized) and Meganthias. Randall & Heemstra 2005. UPRM 3809, 157 mm SL; Mona Passage, (2006: 4) distinguished Odontanthias from Holan- NE of Desecheo Island (Desecheo Island at 18°23’ thias on the basis of the shape of the caudal fin N, 67°28’ W); 188 fathoms (344 meters); 10 (“deeply emarginate with rounded lobes to lunate August 2005. All paratypes collected by Eugenio with slender, sometimes filamentous lobes” in Piñeiro Soler and Miguel A. Vargas. Odontanthias vs. “near-truncate to rounded or Diagnosis: A species of Odontanthias distinguish- rhomboid... with a long slender lobe in the ventral able from other members of the genus by the fol- part of the fin of one of the species” in Holanthias) lowing combination of characters. First three soft and the absence of accessory scales on the body rays of pelvic fin produced, second reaching past scales of species of Odontanthias (although present base of anal fin to well past base of caudal fin. Cau- “on the head and nape of a few species”) vs. dal fin with upper and lower lobes produced into “numerous accessory scales on the body scales of long filaments. Dorsal spines not produced and Holanthias.” without long filamentous projections. No filamen- In the diagnosis presented in the original descrip- tous dorsal and anal soft rays. Vomerine tooth tion of Meganthias, Randall & Heemstra (2006) patch subquadrangular to diamond-shaped, with- wrote that their new genus had the characters of out posterior prolongation. Dorsal fin rays X, 15. Odontanthias except for a number of morphological Pectoral fin rays 18. Lateral line scales 33 to 38. differences, the most important of which would Side of head with two bright yellow stripes; pelvic, seem to be anal soft rays 8 or 9 and the presence of anal and caudal fins bright yellow. accessory scales (“dense on head and nape,” Randall Description: Morphometric data are in Table I. & Heemstra 2006: 27) in Meganthias vs. anal soft Data for countable characters follow; values for the rays 7 or 8 (usually 7) and the absence of accessory holotype are indicated by asterisks for characters scales on the body (“but may be present on head having variable counts (for characters that can be and nape of some species,” p. 4) in Odontanthias. The new species becomes the fourteenth species of Odontanthias to be described, represents the first record of the genus outside of the Indo-Pacific region and is the twenty-third anthiine species recorded from the Atlantic Ocean.

METHODS AND ABBREVIATIONS Methods are those of Anderson & Heemstra Fig. 1. Holotype of Odontanthias hensleyi, USNM 400888, (1980), as modified by Anderson et al. (1990) and 155 mm SL; photographed by D. E. De Vore. aqua vol. 18 no. 1 - 15 January 2012 26 William D. Anderson, Jr. and Graciela García-Moliner

Table I. Morphometric data for Odontanthias hensleyi n. sp. Holotype USNM 400888; Para I = Paratype, UPRM 3809; Para II = Paratype, UPRM 3794; Para III = Paratype, UPRM 3793. Standard lengths in mm, other measurements in percentages of standard length. Dam. = damaged; > = slightly damaged.

Character Holotype Para I Para II Para III Standard length 155 157 159 162 Head, length 36.4 35.4 34.8 35.2 Snout, length 7.4 6.6 6.9 7.6 Bony orbit, diameter 11.6 11.8 11.2 11.0 Postorbital head length 17.3 16.9 15.9 15.5 Upper jaw, length 19.1 17.9 17.7 18.1 Maxilla width 7.1 6.2 7.3 7.1 Lower jaw, length 20.8 18.2 20.0 20.1 Bony interorbital 9.3 8.2 8.7 8.4 Body, depth 41.6 39.0 42.2 40.8 Body, width 16.1 18.4 18.0 17.5 Predorsal-fin length 34.9 34.0 34.1 34.7 Preanal-fin length 63.6 63.7 66.5 67.2 Prepelvic-fin length 37.7 36.7 37.6 37.3 Caudal peduncle, length 24.4 25.5 23.0 24.4 Caudal peduncle, depth 13.8 12.5 13.4 12.8 Dorsal fin, base length 58.9 57.5 60.1 56.1 Anal fin, base length 18.1 16.3 18.5 16.6 Anal fin, length 37.5 34.9 37.7 33.7 Pectoral fin, length 40.4 35.5 43.8 36.1 Pelvic fin, length 79.9 67.8 >65.9 ca. 60.7 Pelvic spine, length 18.2 17.3 17.9 17.0 Upper caudal lobe 85.8 ca. 76.9 >73.4 Dam. Lower caudal lobe 84.5 Dam. >75.3 >65.8 1st dorsal spine 5.9 6.8 5.3 5.8 2nd dorsal spine 9.7 9.1 9.7 8.0 3rd dorsal spine 11.5 10.5 10.7 10.1 4th dorsal spine 13.6 12.8 11.9 10.9 Longest dorsal spine 10th-14.0 9th-13.9 5th-12.2 4th-10.9 1st dorsal soft ray 16.5 15.5 15.5 14.7 Last dorsal soft ray 14.2 14.1 >12.3 14.1 Longest dorsal soft ray 10th-28.4 9th-28.6 10th-27.0 10th-25.0 1st anal spine 7.1 6.1 Dam. 5.4 2nd anal spine 13.0 12.5 11.6 10.8 3rd anal spine 13.5 14.4 Dam. 11.5 Longest anal soft ray 4th-25.9 4th-22.6 4th-21.3 4th-21.5 examined only by radiography, the values are those origin and lateral line 12* to 14. Circumpeduncu- of the holotype). lar scales 16* to 18. Vertebrae 26 (10 precaudal + Dorsal fin rays X, 15. Anal fin rays III, 7. Pectoral 16 caudal). First caudal vertebra without para- fin rays 18. Pelvic fin rays I, 5. Caudal fin rays: pophyses. Formula for configuration of supra- principal 15 (8 + 7); branched 13 (7 + 6); procur- neural bones, anterior neural spines and anterior rent 8 dorsally, 8 ventrally. Branchiostegal rays 7. dorsal pterygiophores 0/0/2/1 + 1/1/ (using nota- Gill rakers on first arch 13* or 14 + 28 to 30 (29*) tion of Ahlstrom et al. 1976). Pleural ribs 8, on (total 42* or 43). Tubed lateral line scales 33* to 38 vertebrae 3 through 10. Epineurals associated with (35*). Rows of cheek scales ca. 7* or 8. Rows of first 11 vertebrae. No trisegmental pterygiophores scales between lateral line and mid-base of spinous associated with dorsal or anal fins. No spur on pos- dorsal fin 1.5* or 2. Scales between dorsal fin ori- teriormost ventral procurrent caudal-fin ray (see gin and lateral line 3 or 4*. Scales between anal fin Johnson 1975); penultimate ventral procurrent

27 aqua vol. 18 no. 1 - 15 January 2012 A n. sp. of Odontanthias Bleeker from Mona Passage off Puerto Rico, the first record of the genus from the Atlantic Ocean caudal fin ray not shortened basally. Parhypural Lateral line complete, anteriorly ascending above with hypurapophysis. Autogenous hypurals 5, no pectoral fin base to run parallel to dorsal body con- hypural fusions. Epurals 3. tour a few scale rows ventral to base of dorsal fin, Mouth terminal, oblique; premaxillae protrusile; then descending precipitously ventral to posterior lower jaw usually exceeding upper jaw slightly with end of soft dorsal fin to run posteriorly near mid- mouth closed. Supramaxilla apparently present dle of caudal peduncle and terminating at distal (due to dense squamation on upper jaw, presence end of hypural bones. Tubes in lateral line scales difficult to determine with certainty). Labial bor- mostly simple, a few scales on holotype and one of der of maxilla with or without hook (see Anderson paratypes (UPRM 3809) with distally branched et al. 1990: 926, fig. 2); holotype: right maxilla tubes. Scales ctenoid with only marginal cteni, i. e., with very short hook, left without hook; on one no ctenial bases present proximal to marginal cteni paratype (UPRM 3793), hook absent but maxilla (see Hughes 1981; this type of scale called periph- widened on both sides at point where hook would eral ctenoid by Roberts 1993). No secondary squa- occur; on second paratype (UPRM 3794) hook mation on body, but a few accessory scales present present on left side, shelf and short anterior projec- on head. Most of head, including dorsum of snout, tion from shelf present on right side; on third interorbital region, maxilla and dentary covered paratype (UPRM 3809) hook present on each side. with scales. Lips, lateral aspect of snout, lachrymal, Posterior border of maxilla truncate; maxilla reach- gular region, branchiostegals and branchiostegal ing posteriorly to vertical through posterior border membranes without scales. Spinous dorsal fin of pupil to as far as vertical through posterior bor- without scales; soft dorsal and anal fins with scales der of orbit. Anterior and posterior nares close to basally and with columns of scales on some inter- eye and to each other, internarial distance 7 to 13 radial membranes; pectoral and pelvic fins scaly times in snout length; anterior naris in short tube, basally and for some distance out onto fins; most posterior border of tube not reaching posterior of caudal fin heavily covered with scales. Pelvic naris when reflected; posterior naris oval, much axillary scales poorly developed or absent; scales in larger than anterior naris, with long axis of oval ventral midline between pelvic fin bases (inter- running dorsoventrally. Interorbital space convex. pelvic process) well developed. No fleshy papillae on border of orbit. Preopercle Dorsal and anal spines without long filaments. without antrorse spines on horizontal limb; hori- Anal fin truncate posteriorly; second anal spine zontal limb with one to a few serrae or with irreg- more robust than first or third. Pectoral fin nearly ular border; vertical limb with numerous small ser- symmetrical, middle rays longest, reaching posteri- rae; region of angle with larger serrae, or with orly to vertical through middle of base of soft dor- roughened border. Distal margins of interopercle sal fin; dorsalmost pectoral fin ray unbranched, and subopercle smooth, or with few small serrae other pectoral-fin rays branched. Pelvic fin inserted near their junction or in places somewhat rough- beneath pectoral fin base; first three pelvic soft rays ened. Premaxilla with series of conical teeth later- produced, second longest, reaching past base of ally and band of villiform teeth medially; at ante- anal fin to well past base of caudal fin. Caudal fin rior end of jaw one or two canine(s) adjacent to lunate with very long filamentous lobes; upper lobe patch of very small teeth; symphysis edentate. 2.1 to 2.4 times as long as head, lower lobe 1.9 to Dentary with 1 to 3 recurved canine(s) about one- 2.3 times as long as head. fourth to one-third way back from anterior end of Coloration: Description based on color pho- jaw; anterior to recurved canine(s) patch or band of tographs of holotype (except where noted) (Fig. 1). villiform to very small conical teeth; posterior to Head mostly reddish orange dorsally and laterally, recurved canine(s) series or band of small conical yellowish ventrally (one paratype mainly red ante- teeth extending along jaw; exserted canine at ante- rior to orbit, mostly violet posterior to orbit, rior end of jaw; symphysis edentate. Vomer, mostly white ventrally). Head with two wavy palatines, endopterygoids and tongue with small bright yellow stripes; dorsal stripe running from teeth; vomerine tooth patch subquadrangular to near midorbit to posterior margin of opercle, ven- diamond shaped, without posterior prolongation; tral stripe extending from below anterior part of palatine and endopterygoid teeth in longitudinal orbit (from near anterior end of snout in paratype) patches (endopterygoid teeth not seen on two of to base of pectoral fin; short yellow band on upper paratypes, UPRM 3793 & 3809). jaw near its anterior end. Iris of eye with bright yel- aqua vol. 18 no. 1 - 15 January 2012 28 William D. Anderson, Jr. and Graciela García-Moliner low circle of pigment surrounding pupil; periph- Linnaean classification of teleosts based on mono- eral to yellow, circle of red, then semicircle of dull phyletic groups, placing the Serranidae in the purple, then semicircle of red (paratype similar but Order Scorpaeniformes, Suborder Serranoidei. with semicircle of bright blue anteriorly). Dorsally Mona Passage is the only known locality for body mostly reddish orange with numerous yellow another relatively recently described fish species. blotches; laterally alternating horizontal silvery and Symphysanodon mona was described from a single reddish orange (or yellowish) lines ventral to lateral specimen collected by the R/V Oregon in October line; ventrally pale; caudal peduncle mostly yellow. 1959, at 18°13’ N, 67°20’ W, in 384 meters Dorsal fin yellow green suffused with red (red ante- (Anderson & Springer 2005). The apparent riorly, blue posteriorly on paratype); pectoral fin restricted distributions of these two species are dull orange (not seen clearly on paratype); pelvic, probably collecting artifacts. anal and caudal fins bright yellow. Etymology: The specific epithet hensleyi is for Comparisons: Based on data and illustrations Dannie A. Hensley, formerly an ichthyologist at provided by Randall & Heemstra (2006), O. hens- the University of Puerto Rico, Mayagüez (Ballan- leyi differs from all but two of the described species tine et al. 2008 and Courtenay et al. 2011 provided of Odontanthias in having 15 soft rays in the dor- obituaries for Hensley, 1944-2008). sal fin, whereas six described species have 14 or The common name Euripos Jewelfish is derived fewer, and five have 16 or more. It differs from the from a Greek word meaning “canal, channel, ditch, other two described species of the genus in having or strait” in allusion to the type locality, Mona Pas- 18 pectoral fin rays, whereas those two have 17 or sage, and the fact that the new species like most fewer. Also, it differs from most other Odontanthias anthiine fishes is a jewel to behold. in having the vomerine teeth in a subquadrangular to diamond-shaped patch without a posterior pro- ADDENDUM longation, in contrast to a variety of shapes in those While the manuscript for this paper was in press, other species (see Randall & Heemstra 2006: 8, fig. the description of another new species of Odontan- 1). In addition O. hensleyi can be distinguished thias was published. White (2011) described Odon- from other Odontanthias by the following combi- tanthias randalli from 11 specimens obtained from nation of characters: pelvic fin reaching past base of a fish market in eastern Lombok, Indonesia. anal fin to well past base of caudal fin; upper and White’s new species can be distinguished from O. lower lobes of caudal fin produced into long fila- hensleyi by the following (data for O. randalli pre- ments; head with two bright yellow stripes on side; cede those for O. hensleyi): soft rays in the dorsal and pelvic, anal and caudal fins bright yellow fin 16 or 17 vs. 15; pectoral-fin rays 15 or 16 vs. (compare Fig. 1, herein, with figs 4 & 5 and plates 18; vomerine tooth patch arrowhead shaped vs. I-VI in Randall & Heemstra 2006). subquadrangular to diamond shaped without pos- Sexuality: Histological examination of gonadal terior prolongation; pelvic, anal and caudal fins tissue shows the holotype of O. hensleyi to be a variously colored vs. bright yellow. With the addi- male. Where investigated histologically and/or tions of O. hensleyi and O. randalli, the number of behaviorally many species of anthiines have been described species in the genus Odontanthias found to be protogynous. Consequently, it would becomes 15. not be surprising to learn that this species is pro- togynous. ACKNOWLEDGEMENTS Distribution: All four specimens of O. hensleyi Captain Eugenio Piñeiro Soler and Miguel A. were collected in Mona Passage, the strait between Vargas collected the specimens of the new species, Puerto Rico and the Dominican Republic leading while fishing commercially for Etelis oculatus from the open Atlantic to the Caribbean Sea. (Queen Snapper) off Rincón on the west coast of Remarks: Due to the unsettled state of the higher Puerto Rico. Matthew T. Craig sent all of the classification of teleosts, we have taken a conserva- known specimens of the new species to the first tive approach and retained the Serranidae in the author. Denise E. De Vore photographed the holo- Order Perciformes, but acknowledge the fact that type and one of the paratypes; Antony S. Harold not all ichthyologists would agree with that assign- produced the radiograph studied; William A. ment. For example, Wiley & Johnson (2010), Roumillat examined and interpreted histological using morphological synapomorphies, proposed a sections of gonadal tissue; and Phillip C. Heemstra

29 aqua vol. 18 no. 1 - 15 January 2012 A n. sp. of Odontanthias Bleeker from Mona Passage off Puerto Rico, the first record of the genus from the Atlantic Ocean and G. David Johnson commented on a presub- 2008. In memoriam: Dannie A. Hensley (1944-2008). mission draft of the manuscript. This is Contribu- Caribbean Journal of Science 44 (2): 273-276. tion Number 369 of the Grice Marine Biological BLEEKER, P. 1873. Sur les espèces indo-archipélagiques Laboratory, College of Charleston. d’Odontanthias et de Pseudopriacanthus. Nederlandsch Tijdschrift voor de Dierkunde 4: 235-240. COURTENAY, W. R., JR., MUNROE, T. A., WINTERBOTTOM, REFERENCES R., RUIZ-CARUS, R. & SMITH-VANIZ, W. F. 2011. Dannie AHLSTROM, E. H., BUTLER, J. L. & SUMIDA, B. Y. 1976. Alan Hensley (1944-2008). Copeia 2011(2): 327-331. Pelagic stromateoid fishes (Pisces, Perciformes) of the HUGHES, D. R. 1981. Development and organization of eastern Pacific: Kinds, distributions, and early life histo- the posterior field of ctenoid scales in the Platycephali- ries and observations on five of those from the northwest dae. Copeia 1981 (3): 596-606. Atlantic. Bulletin of Marine Science 26 (3): 285-402. JOHNSON, G. D. 1975. The procurrent spur: An unde- ANDERSON, W. D., JR. & BALDWIN, C. C. 2000. A new scribed perciform caudal character and its phylogenetic species of Anthias (Teleostei: Serranidae: Anthiinae) from implications. Occasional Papers of the California Academy the Galápagos Islands, with keys to Anthias and eastern of Sciences No. 121: 1-23. Pacific Anthiinae. Proceedings of the Biological Society of RANDALL, J. E. & HEEMSTRA, P. C. 2006. Review of the Washington 113 (2): 369-385. Indo-Pacific fishes of the genus Odontanthias (Ser- ANDERSON, W. D., JR. & HEEMSTRA, P. C. 1980. Two new ranidae: Anthiinae), with descriptions of two new species species of western Atlantic Anthias (Pisces: Serranidae), and a related new genus. Indo-Pacific Fishes 38: 1-32, redescription of A. asperilinguis and review of Holanthias plates I-VIII. martinicensis. Copeia 1980 (1): 72-87. ROBERTS, C. D. 1993. Comparative morphology of spined ANDERSON, W. D., JR., PARIN, N. V. & RANDALL, J. E. scales and their phylogenetic significance in the Teleostei. 1990. A new genus and species of anthiine fish (Pisces: Bulletin of Marine Science 52 (1): 60-113. Serranidae) from the eastern South Pacific with com- WHITE, W. T. 2011. Odontanthias randalli n. sp., a new ments on anthiine relationships. Proceedings of the Biolog- anthiine fish (Serranidae: Anthiinae) from Indonesia. ical Society of Washington 103 (4): 922-930. Zootaxa 3015: 21-28. ANDERSON, W. D., JR. & SPRINGER, V. G. 2005. Review of WILEY, E. O. & JOHNSON, G. D. 2010. A teleost classifi- the perciform fish genus Symphysanodon Bleeker (Sym- cation based on monophyletic groups. Pp. 123-182. In: physanodontidae), with descriptions of three new species, Origin and phylogenetic interrelationships of teleosts. Eds S. mona, S. parini, and S. rhax. Zootaxa 996: 1-44. Nelson, J. S., Schultze, H.-P. and Wilson, M. V. H. Dr. BALLANTINE, D. L., APPELDOORN, R. S. & APONTE, N. E. Friedrich Pfeil, München.

aqua vol. 18 no. 1 - 15 January 2012 30 aqua, International Journal of Ichthyology

Threatened fishes of the world: Stenodus leucichthys leucichthys Güldenstädt, 1772 (Salmonidae)

Samaneh Poursaeid & Bahram Falahatkar*

Fisheries Department, Faculty of Natural Resources, University of Guilan, Sowmeh Sara, Guilan, Iran * Corresponding author. Fisheries Dept., Faculty of Natural Resources, University of Guilan, Sowmeh Sara, P.O. Box 1144, Iran. Tel.: +98 182 322 3599; Fax: +98 182 322 2102. E-mail: [email protected]

Received: 24 April 2011 – Accepted: 30 July 2011

Abstract mique dans la région,mais il y a peu de données concernat This paper introduces the ecological and biological char- les populations de la mer Caspienne. Nous traitons de sa di- acteristics of the Inconnu (Stenodus leucichthys leucichthys). stribution, de ses caractéristiques écologiques et reproduc- This is a Caspian Sea endemic species of the family tives et des causes de son extinction en mer Caspienne. Salmonidae, listed as Extinct in the Wild in IUCN’s Red Data Book due to restricted access to spawning grounds, Sommario dam construction, illegal fishing, and environmental pol- Questo articolo presenta le caratteristiche ecologiche e bio- lution. This valuable species has considerable ecological logiche del salmone bianco noto anche come “Inconnu” and economical importance for the region, but there are (Stenodus leucichthys leucichthys). Si tratta di una specie ende - little data for the Caspian Sea populations. We discuss its mica del Mar Caspio appartenente alla famiglia Salmonidae distribution, ecological and reproductive characteristics, e elencata come “Estinta in natura” dallo IUCN Red Data and the causes of its extinction in the Caspian Sea. Book a causa delle limitazioni degli spazi riproduttivi, della costruzione di dighe, della pesca illegale e dell’inquinamento Zusammenfassung ambientale. Questa specie ha un’importanza considerevole In dieser Arbeit werden die ökologischen und biologischen per la regione dal punto di vista ecologico ed economico, ma Merkmale des Weißlachses („Inconnu“) Stenodus leucichthys esistono pochi dati per le popolazioni del Mar Caspio. Se ne leucichthys einführend behandelt. Es handelt sich um eine discutono la distribuzione, l’ecologia, la riproduzione e le endemische Art des Kaspischen Meeres aus der Familie der cause dell’estinzione nel Mar Caspio. Lachsfische (Salmonidae), die in der Roten Liste des IUCN als „ausgestorben“ geführt wird; als Gründe werden Ein- INTRODUCTION schränkung der Laichgründe, Staudammbauten, illegale The Inconnu (Stenodus leucichthys leucichthys Befischung und Umweltverschmutzung genannt. Die Güldenstädt) is the largest and fastest-growing wertvolle Art hat erhebliche ökologische und ökonomische Bedeutung für die Region, aber es gibt wenige Daten über member of the whitefish subfamily Coregoninae die Populationen des Kaspischen Meeres. Wir diskutieren (Salmonidae). Two subspecies of Inconnu are rec- die Verbreitung, die ökologischen und fortpflanzungsbiolo- ognized, with S. l. leucichthys isolated in the gischen Kennzeichen sowie die Gründe für die Ausrottung Caspian Sea drainage of western Asia, while S. l. im Kaspischen Meer. nelma (Pallas) is found in the Arctic and sub-arctic regions of north-western North America and north- Résumé ern Eurasia (Stephenson et al. 2005). It is known by Cet article propose les caractéristiques écologiques et a range of common names: Inconnu (English), biologiques du Stenodus (Stenodus leucichthys leucichthys). Il Belorybitsa (Russian), Mahi Ziba (Persian), Stenode s’agit d’une espèce endémique de la Caspienne, de la famille de Salmonidae, répertoriée comme étiente dans la nature Blanc (French), Salmon Blanco (Spanish) Ak balyk selon l’IUCN’s Red Data Book, à cause de l’accès ma laisé (Kazakh) and Azatmahy (Turkmenian). aux frayères, à la construction d’un barrage, à la pê che illé- Stenodus leucichthys leucichthys is the Caspian gale et à la pollution de l’environnement. Cette espè ce a une endemic subspecies and mainly occurs along the importance considérable sur le plan éco logi que et écono - western and eastern coasts of the middle Caspian

31 aqua vol. 18 no. 1 - 15 January 2012 Threatened fishes of the world: Stenodus leucichthys leucichthys Güldenstädt, 1772 (Salmonidae)

Sea at depths of 60-65 m (Berg 1948). Declining imum length and weight 130 cm and 35 kg respec- natural populations in the Caspian Sea due to tively; much larger individuals have been recorded. heavy fishing pressure, increasing water pollution Sides of body silvery, belly silvery white without and habitat alteration and destruction have spots, dorsum usually green, blue or pale brown resulted in S. l .leucichthys being listed as “Extinct (Fig. 1). Sexual dimorphism develops during in the wild” (IUCN Red List 2010). Concerns spawning period, when epithelial tubercles appear about the decline of Stenodus l. leucichthys popula- on head and side of body of males (Berg 1948; tions led Volga River hatcheries to carry out artifi- Shariaty 2001). cial reproduction and restocking in the Caspian Distribution: Stenodus l. leucichthys is found only Sea (Belyaeva & Milstein 1959). The Arctic sub- in rivers draining to the Caspian Sea, from which it species, Stenodus leucichthys nelma, was introduced mainly ascends the Volga, while very few fish into the northern Volga River drainage and is now ascend the Ural, and it is rarely found in the Terek. expanding, so may threaten surviving populations It is widely found along the western coast north of and cultivated stocks through hybridization (Frey- Makhach-kala and along the eastern coast hof & Kottelat 2008). (Mangyshlak). Its closest relative, Stenodus leuci- Despite the ecological and economic importance chthys nelma, penetrates into the Caspian Sea from of this fish, very little is known about it in the the Arctic Ocean basin through the Post-Glacial Caspian Sea. Therefore, the aim of this study is to ponded lakes which are situated between the basins introduce and better understand the ecological and of the Volga and the Kama, on the one hand, and biological characteristics of this valuable species. of the Baltic Sea and the Arctic Ocean, on the other Identification: D II-VI (9-13), A II-IV 9-15, dor- (Berg 1948). Stenodus l. leucichthys spends the warm sal fin high and pointed, adipose fin present, pelvic season in the central and southern regions of the fins with well developed axillary process, vertebrae Caspian Sea (Kottelat & Freyhof 2007) mainly in 65-68, manubrium absent, with hook-shaped Guilan and no record of this species has been processes on the capitulum, anterior edge of reported from the Mazandaran and Golestan coasts hyomandibular round, only 19 to 26 gill rakers (FAO 1991). Figure 2 shows the distribution of S. present on lower limb of first gill arch. Scales large, l. leucichthys in the Caspian Sea basin. 99-120 in lateral line. Body fusiform and moder- Habitat and ecology: Stenodus l. leucichthys is a ately slender, head relatively small, mouth large pelagic species which inhabits open waters to the and terminal, lower jaw long, tip projecting, hind depth 65 m, and has not been found below 65m end reaching back behind eye, upper jaw reaching (its optimum depths are 25-45 m). This fish is an to level of pupil, teeth present on jaws, vomer, oxyphilic species and prefers waters with tempera- palatines and tongue. Pyloric caeca 191-193. Max- ture below 20°C. It is heterotrophic and an active

Fig. 1. Stenodus leucichthys leucichthys from the southern part of the Caspian Sea; weight 4300 g; total length 75 cm. Photo by S. Poursaeid. aqua vol. 18 no. 1 - 15 January 2012 32 Samaneh Poursaeid & Bahram Falahatkar predator. At 30 days after hatching, fingerlings BRIEF DISCUSSION begin to feed on invertebrates and larvae and small Unfortunately, the published data on different juveniles of other fishes. In the sea, adult S. l. leuci- aspects of S. l. leucichthys is limited and most stud- chthys feeds mainly on small fish (clupeids, ies on the species has been carried out on North engraulids, juvenile cyprinids, atherinids, gobiids American and Eurasian populations of Stenodus etc) (Podlesny 1947; Svetovidov 1984; Shariaty leucichthys nelma (Scott & Crossman 1973; Alt 2001). Stenodus l. leucichthys is amphidromous. 1977; Morrow 1980; Brown 2000; Howland et al. However, amphidromy is not obligatory and non- 2000; Underwood et al. 2000; Stephenson et al. amphidromous populations have been described in 2005). Therefore, much effort from scientists and some locations (Petrova 1976). Mature adults migrate from sea to the delta of the Volga in the fall, winter and the early spring. Therefore, the species has two seasonal forms, spring and autumn, which are differentiated in time of entry to the Volga for spawning (Berg 1948). Adults migrate downward to the sea after spawning, but many of them die (Freyhof & Kottelat 2008). The alevins immedi- ately descend to the sea after hatching. Reproduction: The main spawning sites of S. l. leucichthys are located in the basin of the Kama, in the Ufa River, 3000 km upstream from the mouth of the Volga (Shariaty 2001). The secondary spawn- ing grounds are situated in the Belaya between Ufa and Sterlitamak. The spawning season in the Ufa begins from the middle of October to the begin- ning of November. Males mature at +5 to +6 years while females reach maturity at the age of +6 to +7 years (Freyhof & Kottelat 2008). Stenodus l. leuci- chthys spawns twice during its life cycle with an interval of two years (Berg 1948). Its optimum tem- perature for reproduction is 0.1 to 6°C. Semi-adhe- sive eggs are deposited on the gravel and rock sub- strate (Kottelat & Freyhof 2007). The sex ratio of brooders in the natural spawning grounds is almost 1:1 as for the subspecies Stenodus leucichthys nelma (Brown 2000). The average fecundity is 250× 103 eggs per individual (104.5-400× 103). On average, about 26% of the total weight of each fish is egg weight (Berg 1948). Embryonic development takes about 180-200 days. Fry hatch from March up to the early May (mostly the second half of April). From the seventh day onward, fry begin to feed on plankton. The fry stage lasts around two months and then they develop into fingerlings (Berg 1948). Threats: The sharp decline in their abundance due to the construction of dams, insufficient spawning areas, increasing illegal fishing, unstable hydrological conditions, river contamination and damage caused by other fish (especially kilka) and crustaceans has resulted in this species being listed as extinct in the wild (Letichevski 1983; IUCN Fig. 2. Distribution of Stenodus leucichthys leucichthys in 2010). the Caspian Sea basin (red circles).

33 aqua vol. 18 no. 1 - 15 January 2012 Threatened fishes of the world: Stenodus leucichthys leucichthys Güldenstädt, 1772 (Salmonidae) governments should be made in understanding the BROWN, R. J. 2000. Migratory patterns of Yukon River biology, ecology and behaviour for managing this Inconnu as determined with otolith microchemistry and valuable species. One key objective in working radio telemetry. MSc thesis. Fairbanks, Alaska, 73 pp. with endangered species is to increase the number FAO. 1991. New species of freshwater fish from Iranian coast of the Caspian Sea. FI: UNDP/IRA/88/001, Rome, of individuals of the species concerned by artifi- Italy. cially/controlled reproduction in captivity. Unfor- FREYHOF, J. & KOTTELAT, M. 2008. Stenodus leucichthys. tunately, there is no policy for preserving this In: IUCN 2010. IUCN Red List of Threatened Species. species in the Caspian Sea from illegal fishermen Version 2010.4. . Downloaded on by countries bordering the Sea and this should be 31 January 2011. done according to fishing methods, size of net HOWLAND, K. L., TALLMAN, R. F. & TONN, W. M. 2000. mesh and time/place prohibition. Previously, the Migration patterns of freshwater and anadromous Russian federation enhanced the spawning areas in Inconnu in the Mackenzie River system. Transactions of the lower parts of the Volgograd hydroelectric the American Fisheries Society 129: 41-59. IUCN. 2010. IUCN Red List of Threatened Species. power station for increased efficiency of natural www.iucnredlist.org. Downloaded on 31 January 2011. reproduction. KOTTELAT, M. & FREYHOF, J. 2007. Handbook of Euro- In connection with the present study, the negative pean Freshwater Fishes. Publications Kottelat, Cornol, influences of the modern environment needs to be Switzerland: 646 pp. studied (i.e. all possible causes of stress, environ- LETICHEVSKI, M. A. 1983. Reproduction of Inconnu. mental changes, destruction of spawning grounds) Moscow. “Lyogkaya I pischevaya promyshlennost” (Light in order to better organize protection of this species and Food Industry): 30-112. and thus develop the necessary strategies for action. MORROW, J. E. 1980. The Freshwater Fishes of Alaska. Findings from this preliminary data suggest that Alaska Northwest Publishing Company, Anchorage, Alaska, 248 pp. further research is needed for the future, including PETROVA, N. A. 1976. The biology of the Inconnu, Sten- population genetics, artificial spawning, domesti- odus leucichthys nelma, from the Irtysh River basin. Jour- cation for future aquaculture activities, natural nal of Ichthyology 16: 17-27. behaviour during migration and spawning, as well PODLESNY, A. V. 1947. Inconnu Stenodus leucichthys Güld. as ionic balance and osmoregulation. The collec- Bio-ecological outline essay. Proc. Siberian Department tive goal of this research should enhance the effec- VNIORKh (All-Union Research Institute of Lake and tiveness of breeding programs, increase popula- Riverine Fisheries) 7: 1:3-195. tions of remaining wild stock, and to improve our SCOTT, W. B. & CROSSMAN, E. J. 1973. Freshwater Fishes of Canada. Fisheries Research Board of Canada, Bulletin understanding of the biological knowledge of this 184, Ottawa, Ontario. 966 pp. species. Since stocks have declined, much effort is SHARIATY, A. 2001. Breeding and rearing Stenodus leuci- needed to rehabilitate the wild populations by all chthys. Shahr Sabz Publication, Rasht, Iran, 82 pp. countries around the Caspian Sea. STEPHENSON, S. A., BURROWS, J. A. & BABALUK, J. A. 2005. Long-distance migrations by Inconnu (Stenodus REFERENCES leucichthys) in the Mackenzie River System. Arctic 58: ALT, K. T. 1977. Inconnu, Stenodus leucichthys, migration 21-25. studies in Alaska 1961-1977. Journal of the Fisheries SVETOVIDOV, A. N. 1984. Salmonidae. In: Fishes of the Research Board of Canada 34: 129-133. north-eastern Atlantic and the Mediterranean 1. (Eds P. J. BELYAEVA, V. N. & MILSTEIN, V. V. 1959. Rearing of P. W HITEHEAD, M. L. BAUCHOT, J. C. HUREAU, J. Inconnu fingerlings in the Volga river delta. Moscow: NIELSEN & E. TORTONESE.): 373-385. “Rybnoye Khozyaistvo”. Fish Industry Journal: 1-18. UNDERWOOD, T. J. 2000. Abundance, length composi- BERG, L. S. 1948. Fishes of freshwaters of USSR and adja- tion, and migration of spawning inconnu in the Selawik cent countries. Vol I. USSR Academy of Sciences pub- River, Alaska. North American Journal of Fisheries Man- lishers, Moscow-Leningrad. agement 20: 386-393.

aqua vol. 18 no. 1 - 15 January 2012 34 aqua, International Journal of Ichthyology

Description of a new Snake Eel (Pisces: Ophichthidae: Myrichthys) from the Philippines

John E. McCosker1 and Gerald R. Allen2

1) California Academy of Sciences, San Francisco, CA 94118, USA. E-mail: [email protected] 2) Western Australian Museum, Locked Bag 49, Welshpool DC, Perth, Western Australia 6986. E-mail: [email protected]

Received: 13 September 2011 – Accepted 10 October 2011

Abstract Vie ne fornita una chiave per l’identificazione delle specie Myrichthys paleracio new species is described from two di Myrichthys. specimens collected in shallow-water coral reefs from the Verde Passage, southern Luzon Island, Philippines. It dif- fers from all known Myrichthys in its vivid brown-and- INTRODUCTION white coloration, its body elongation (body depth 43 times The snake eels of the cosmopolitan genus in total length) and its mean vertebral formula Myrichthys were revised by McCosker & Rosen- (3/77.5/183). A key to the species of Myrichthys is pro- blatt (1993). They recognized nine valid species vided. and 28 synonyms. We are unaware of subsequent publications that have either described additional Zusammenfassung species or changed the taxonomy of the taxa Die neue Art Myrichthys paleracio wird auf der Grundlage treated therein. We herein describe a tenth species von zwei Exemplaren beschrieben, die über Korallenriffen from the Verde Passage, southern Luzon Island, im Flachwasser in der Verde-Passage, südlich der Insel Philippines. Two specimens of this vividly pat- Luzon, Philippinen, gefangen worden waren. Sie unter- terned snake eel were collected by Peri Paleracio, an scheidet sich von allen anderen Myrichthys-Arten durch die avid naturalist, diver, and fish collector, and we are lebhaft braun-weiße Farbgebung, den längeren Körper pleased to name it in his honor. (Gesamtlänge gleich 43-fache Körpertiefe) und die mit- tlere Wirbelformel (3/77,5/183). Außerdem wird ein Be- MATERIALS AND METHODS stimmungsschlüssel der Myrichthys-Arten wiedergegeben. Measurements are straight-line, made either with a 300 mm ruler with 0.5 mm gradations (for total Résumé length, trunk length, and tail length) and recorded Myrichthys paleracio, nouvelle espèce, est décrit sur base to the nearest 0.5 mm, or with dial calipers (all de deux spécimens collectés dans des récifs de corail peu profonds de Verde Passage, région sud de l’île de Luzon, other measurements) and recorded to the nearest Philippines. Il se distingue de tous les Myrichthys connus 0.1 mm. Body length comprises head and trunk par sa colortion brune et blanche, sa forme allongée lengths. Head length is measured from the snout (longueur totale à raison de 43 fois la hauteur du corps) et tip to the posterodorsal margin of the gill opening; sa formule vertébrale principale (3/77,5/183). Une clé trunk length is taken from the end of the head to pour les espèces de Myrichthys est fournie. mid-anus; and maximum body depth does not include the median fins. Head-pore terminology Sommario follows that of McCosker et al. (1989: 257), such La nuova specie Myrichthys paleracio è descritta sulla base that the supraorbital pores are expressed as the eth- di due esemplari raccolti in acque poco profonde lungo la barriera corallina di Verde Passage, isola di Luzon meridi- moidal pore + pores in supraorbital canal, i.e., 1 + onale, Filippine. Essa differisce dagli altri membri del 3, and the infraorbital pores are expressed as pores genere Myrichthys per la vivace colorazione marrone e bian- along the upper jaw + those in vertical part of canal ca, il corpo più allungato (l’altezza sta 43 volte nella lun- behind eye (the “postorbital pores”), i.e., 4 + 2, in ghez za totale) e per la formula vertebrale (3/77.5/183). that frequently the last pore included along the

35 aqua vol. 18 no. 1 - 15 January 2012 Description of a new Snake Eel (Pisces: Ophichthidae: Myrichthys) from the Philippines upper jaw is part of the postorbital series. Vertebral posterior nostrils in upper lip, not visible exter- counts (which include the hypural) were taken nally, beginning before eye and ending beneath from radiographs. The mean vertebral formula middle of pupil; upper lip papillate, particularly (MVF) is expressed as the average of predorsal, pre- between anterior and posterior nostrils; broad anal, and total vertebrae (Böhlke 1982). Institu- fleshy chevron dividing snout between anterior tional abbreviations follow the Standard Symbolic nostrils; dorsal fin origin on head, well in advance Codes for Institutional Research Collections in of gill opening, 1.5-1.7 in HL; pectoral fin minute, Herpetology and Ichthyology (Leviton et al. much shorter than its base, nearly twice in eye 1985). diameter. Head pores minute, but typical of Myrichthys (McCosker 1977); five mandibular, two preoper- Myrichthys paleracio, n. sp. cular, single ethmoidal + 3 supraorbital, 4 + 2 Peri’s snake eel (Figs 1-3) infraorbital, three temporal and single interorbital and supratemporal pores; preopercular, temporal, Holotype: CAS 233313, 311 mm, an immature suborbital, postorbital and other series present; sin- male, Layag Layag, 13.688o N 120.841o E, Luzon gle median interorbital and temporal pores; two Island, Batangas Province, Philippines, hand net, P. preopercular pores; four mandibular pores; lateral Paleracio, 15 May 2011. line pores present, 10 before the gill openings, Paratype: WAM 33154.001, 315+ mm (tail bro- remainder too difficult to ascertain. ken and healed), ripe female, Caban Island, Teeth granular, small and fixed, irregularly biser- 13°41.376’ N 120°50.374’ E), Verde Passage, ial in jaws and on vomer; small intermaxillary Batangas Province, Philippines, mixed sand, rubble chevron anteriorly, followed by gap. and coral, 33m, hand net, P. Paleracio, 12 June Colour in life (Figs 1-3) and in preser- 2009. v a t i o n : white to pale, overlain on head, trunk Diagnosis: An elongate species of Myrichthys with and tail with about 50 or more brown saddles, depth 43 times and tail 1.8 in TL; pectoral fin extending from base of dorsal fin to ventral edge of minute, its length about twice in eye; numerous flank (many saddles incomplete, some irregular, large brown spots on head and body; and total ver- and all wider than pale interspaces, meeting along tebrae 183, mean vertebral formula 3/77.5/183. ventral surface posterior to anus); snout, chin, and Counts and measurements (in mm) of the holo- anterior nostrils markedly white; eye within brown type (followed by those of the paratype in paren- mask, followed by two diagonal bands and several theses): Total length 311 (315+, damaged tail); eye-sized brown spots; throat and anteroventral head 23.7 (35.3); trunk 120.3 (186.7); tail 167 (?); trunk region overlain with several eye-sized brown predorsal distance 13.6 (23.6); pectoral fin length spots; pectoral fins pale; median fin margins pale, 1.2 (2.1); pectoral fin base 2.2 (4.4); body depth at brown body saddles extending onto base of dorsal gill openings ~7.2 (~16); body width at gill open- fins; tail tip pale. ings ~5.5 (~10.5); snout 4.2 (6.9); tip of snout to Remarks: The new species differs from its con- rictus of jaw 6.6 (10.5); tip of chin to rictus of jaw geners on the basis of its coloration, reduced pec- 5.0 (7.9); eye diameter 2.2 (3.9); interorbital dis- toral fin and vertebral number. It is most closely tance 3.6 (6.6); gill opening height 2.1 (4.0); isth- related to the dark-spotted and banded Indo- mus width ~3.8 (~7.0). Predorsal vertebrae 2 (4), Pacific species Myrichthys maculosus (Fig. 3) and M. preanal vertebrae 79 (76); total vertebrae 183. Ten colubrinus and differs from them in the distribu- lateral line pores in left branchial region; remainder tion, number and size of its spots and in its verte- small and difficult to discern. bral number. Myrichthys maculosus (Fig. 3) and M. Description: Body elongate, its depth at gill colubrinus also occur in shallow water within the openings 43 in TL; head and trunk 2.16 and head Philippines Archipelago, however none were 13.1 in TL; snout rounded, conical when viewed observed at the locations where M. paleracio was from above; lower jaw included, snout tip reaching captured. Other individuals were seen but not col- base of anterior nostrils; eye large, 2.7-3.0 in upper lected from Layag Layag. jaw, its center well behind midpoint of upper jaw; The paratype, although a larger specimen than anterior nostrils tubular, elongate, about twice in the holotype, has had much of its tail bitten off and eye, with small lappet extending from each side; regrown (we calculate that it may have been aqua vol. 18 no. 1 - 15 January 2012 36 John E. McCosker and Gerald R. Allen

Fig. 1. Myrichthys paleracio, approximately 500 mm TL, underwater photograph, Caban Island, Verde Channel, Luzon, Philippines. Photo by G. R. Allen.

Fig. 2. Myrichthys paleracio, paratype, 315 mm TL (tail damaged), underwater photograph, Caban Island, Verde Channel, Luzon, Philippines. Photo by G. R. Allen.

37 aqua vol. 18 no. 1 - 15 January 2012 Description of a new Snake Eel (Pisces: Ophichthidae: Myrichthys) from the Philippines

Fig. 3. Comparison of head coloration of Myrichthys (from top): M. paleracio. Two photos by G. R. Allen. M. maculosus. Photo by R. Steene at Miyake-jima, Japan. M. colubrinus. Photo by R. Myers at Fiji. aqua vol. 18 no. 1 - 15 January 2012 38 John E. McCosker and Gerald R. Allen approximately 480mm in length). We have there- 8th vertebra, 1.1-1.4 in head length; total ver- fore selected the smaller but complete specimen to tebrae 159-167...... be the holotype. M. aspetocheiros McCosker & Rosenblatt (eastern Pacific). Key to the species of Myrichthys (modified from 7a. Total vertebrae 177-197...... 8 McCosker and Rosenblatt 1993) 7b. Total vertebrae 149-168...... 9 8a. Total vertebrae 177-183; spotting on chin and 1a. Body coloration pale, overlain with 25-55 throat of adults smaller than or equal to eye, black or brown rings or saddles, separated by spotting on flanks round...... white or pale interspaces, encircling or par- M. magnificus (Abbott) (Hawaii, Leeward and tially encircling the body; body very elongate, Johnston islands). its depth 43-70 times in total length...... 2 8b. Total vertebrae 180-197; spotting on chin and 1b. Body coloration pale or dark, overlain with throat of adults larger than eye, spotting on round spots which are either dark, pale, or flanks generally ovoid...... dark diffuse spots with bright centers; anal and ...... M. maculosus (Cuvier) (Indo-Pacific) dorsal fin end about equally relative to tail tip; 9a. Total vertebrae 149-156 ...... body moderately elongate, its depth 25-45 ...... M. tigrinus Girard (eastern Pacific) times in total length...... 3 9b. Total vertebrae 158-168 ...... 2a. Anal fin ends about 2 head lengths before tail M. pantostigmius Jordan & McGregor (Revil- tip, well in advance of end of dorsal fin; body lagigedo and Clipperton islands). overlain with 25-35 black or brown rings or saddles, separated by white or pale interspaces Behaviour and ecology: The new species was which are equal to or wider than the darker observed underwater with the use of scuba gear by rings; body extremely elongate, its depth 50- the second author at Caban Island (capture site of 70 times in total length...... paratype). Three individuals were seen at depths of ...... M. colubrinus (Boddaert) (Indo-Pacific) 25-35 m on a relatively flat, low relief bottom with 2b. Anal fin ends less than a head length before mixed sand, rubble and small coral formations. The tail tip, at the same level as the end of the dor- eels were invariably encountered in the open and sal fin; body overlain with 50-55 brown rings were swimming slowly, apparently searching the bot- and saddles, separated by white interspaces tom for prey (probably small fishes or crustaceans). which are narrower than the darker rings; Distribution: Myrichthys paleracio is currently body depth 43 times in total length ...... known only from the Verde Channel area of south- ...... M. paleracio n. sp. (Philippines) ern Luzon, Batangas Province, Philippines in 25- 3a. Body coloration of round pale spots on a dark 35 m. background of brown or green ...... Etymology: We take pleasure in naming this new M. breviceps (Richardson) (western Atlantic) eel in honor of its collector, Peri Paleracio, to be 3b. Body coloration pale, overlain with dark spots treated as a noun in apposition. or diffuse dark spots with bright centers ..... 4 4a. Spots on body diffuse with bright centers ACKNOWLEDGEMENTS (gold in life)...... 5 We are grateful to the California Academy of Sci- 4b. Spots on body dark and distinct, without pale ences, which sponsored the first author’s participa- or bright centers...... 6 tion in the 2011 Hearst Expedition to the Philip- 5a. Total vertebrae 151-159 ...... pines. Collection and curatorial assistance for this M. pardalis (Valenciennes) (eastern Atlantic) trip was kindly provided by David Catania (CAS). 5b. Total vertebrae 164-173 ...... We also thank Roger Steene of Cairns, Australia, ...... M. ocellatus (Lesueur) (western Atlantic) for his companionship and diving assistance dur- 6a. Pectoral fin reduced, its length less than the ing the second author’s 2009 trip to the Verde width at its base; dorsal fin origin above 1st, Channel area during which the first specimen was 2nd or 3rd vertebra, 1.4-2.0 in head length; collected. Roger Steene and Robert Myers gener- total vertebrae 149-197...... 7 ously provided underwater photographs. 6b. Pectoral fin developed, longer than the width at its base; dorsal fin origin above 6th, 7th or

39 aqua vol. 18 no. 1 - 15 January 2012 Description of a new Snake Eel (Pisces: Ophichthidae: Myrichthys) from the Philippines

REFERENCES MCCOSKER, J. E., BÖHLKE, E. B. & BÖHLKE, J. E. 1989. BÖHLKE, E. B. 1982. Vertebral formulae of type specimens Family Ophichthidae. In: Fishes of the Western North of eels (Pisces: Anguilliformes). Proceedings of the Aca - Atlantic, Part 9, Volume 1, Böhlke, E. B. (Ed.), pp. 254- demy of Natural Sciences of Philadelphia 134: 31-49. 412. Memoirs of the Sears Foundation for Marine Research, LEVITON, A. E., GIBBS, R. H. JR., HEAL, E. & DAWSON, C. New Haven. E. 1985. Standards in herpetology and ichthyology: part MCCOSKER, J. E. & ROSENBLATT, R. H. 1993. A revision I. Standard symbolic codes for institutional resources col- of the snake eel genus Myrichthys (Anguilliformes: lections in herpetology and ichthyology. Copeia 1985: Ophichthidae) with the description of a new eastern 802-832. Pacific species. Proceedings of the California Academy Sci- MCCOSKER, J. E, 1977. The osteology, classification, and ences 48 (8): 153-169. relationships of the eel family Ophichthidae. Proceedings of the California Academy of Sciences series 4, 41 (1): 1-123.

aqua vol. 18 no. 1 - 15 January 2012 40 aqua, International Journal of Ichthyology

Color variation of the puffer Arothron hispidus (Linnaeus) and comparison with A. reticularis (Bloch & Schneider)

John E. Randall1, Sergey V. Bogorodsky2 and Jean Michel Rose3

1) Bishop Museum, 1525 Bernice St., Honolulu, HI 96817-2704, USA. E-mail: [email protected] 2) Station of Naturalists, Omsk, Russia. E-mail: [email protected] 3) Institut H.E. Sauvage, 56, rue du Dr. Brousse, 62360 St., Etienne-au-Mont, France. E-mail: [email protected]

Received: 13 August 2011 – Accepted 10 October 2011

Abstract Exemplare der verschiedenen Farbformen, insbesondere The Whitespotted Puffer, Arothron hispidus (Linnaeus), vom Roten Meer, zu beschaffen, um die morphologischen wide-ranging in the Indo-Pacific region and the tropical Merkmale, einen möglichen farblichen Geschlechtsunter- eastern Pacific, is very variable in color pattern. It has been schied (Dichromatismus) und molekulare Kennzeichen misidentified as A. reticularis (Bloch & Schneider), which genauer untersuchen zu können. is also Indo-Pacific in distribution. Color variation of A. hispidus is documented, particularly in the Red Sea where Résumé it is unusually diverse in color pattern. The record of A. Arothron hispidus (Linné), largement répandu dans reticularis from the southern Red Sea by Roux-Estève l’Indo-Pacifique et l’est tropical du Pacifique, est de colora- (1956), recognized in three checklists of Red Sea fishes, is tion très variable. Il a été identifié erronément comme A. invalidated. The two species are distinguished by the area reticularis (Bloch & Schneider) dont la distribution con- of skin covered by spinules and by color pattern. A cerne aussi l’Indo-Pacifique. La variation de couleur d’A. subadult specimen of A. reticularis collected from fresh hispidus est documentée, particulièrement en mer Rouge water in Palau represents a first record for Micronesia and où son patron de coloration est inhabituellement diversi- an underwater photograph is a first record for New Cale- fié. La présence d’A. reticularis au sud de la mer Rouge sig- donia. The need is stressed for specimens of the various nalée par Roux-Estève (1956), reprise dans trois relevés de color forms of A. hispidus, especially in the Red Sea, for poissons de la mer Rouge, est invalidée. Les deux espèces morphological study, determination of possible sexual se di stinguent par la zone de peau couverte de spinules et dichromatism and for molecular research. par la coloration. Un spécimen subadulte d’A. reticularis, collecté en eau douce à Palau, est une première occurrence Zusammemfassung pour la Micronésie et une photographie sous-marine con- Der Weißflecken-Kugelfisch Arothron hispidus (Lin- stitue une première pour la Nouvelle Calédonie. Le besoin naeus), der im indo-pazifischen Raum und im tropischen de spécimens de diverses variétés de couleur d’A. hispidus, Ostpazifik weit verbreitet ist, variiiert stark im Farbmuster. surtout en mer Rouge, se fait sentir pour une étude Er lässt sich leicht mit A. reticularis (Bloch & Schneider) morpholo gique, la détermination d’un possible dichroma- verwechseln, der ebenfalls im indopazifischen Raum ver- tisme sex uel et l’analyse moléculaire. breitet ist. Die Farbvariationen von A. hispidus werden dokumentiert, vor allem anhand von Exemplaren vom Sommario Roten Meer, wo es ungewöhnlich stark abweichende Farb- Il pesce riccio, Arothron hispidus (Linnaeus), ampiamente muster gibt. Die Aufzeichnung von A. reticularis vom diffuso nella regione indopacifica e nelle zone tropicali del Roten Meer durch Roux-Estève (1956), die in drei Check- Pacifico orientale, ha una colorazione molto variabile. E’ listen von Fischen des Roten Meeres anerkannt worden stato confuso con A. reticularis (Bloch & Schneider), war, wird für ungültig erklärt. Die beiden Arten lassen sich anch’esso presente nell’Indo-Pacifico. Sono documentate le am Hautbereich, der von Stacheln bedeckt wird, und an varianti di A. hispidus, soprattutto nel Mar Rosso dove der Farbgebung unterscheiden. Ein subadultes Exemplar esiste un’insolita diversità di colorazioni. La segnalazione di von A. reticularis, das im Süßwasser auf Palau gefangen A. reticularis nel Mar Rosso meridionale da parte di Roux- wurde, ist als Erstnachweis für Mikronesien zu werten, Estève (1956), riconosciuta in tre checklist dei pesci del und eine Unterwasserfotografie liefert die erste Aufzeich- Mar Rosso, è invalidata. Le due specie si distinguono per nung für Neukaledonien. Betont wird die Notwendigkeit, l’area di pelle coperta da spinule e per la colorazione. Un

41 aqua vol. 18 no. 1 - 15 January 2012 Color variation of the puffer Arothron hispidus (Linnaeus) and comparison with A. reticularis (Bloch & Schneider) subadulto di A. reticularis raccolto nelle acque dolci di (1993: pl. 205, fig. 7) used an illustration of A. retic- Palau rappresenta la prima segnalazione per la Micronesia ularis to represent A. hispidus. Adding to the confu- e una fotografia subacquea la prima per la Nuova Caledo- sion has been the extreme variation in color exhib- nia. Si sottolinea la necessità di incrementare il numero di ited by what we believe to be one species, Arothron esemplari di A. hispidus di varia colorazione, soprattutto del Mar Rosso, per studi morfologici, per la determi- hispidus, in the Red Sea. The most perplexing are nazione del dicromismo sessuale e per ricerche molecolari. large individuals that have been photographed underwater with either a reticular or a linear pattern INTRODUCTION instead of a pattern of mainly white spots. We The Whitespotted Puffer, Arothron hispidus (Lin- include here a diagnosis of A. hispidus, a comparison naeus, 1758), type locality India, is the most wide- with A. reticularis and photographs of both species. spread of the tetraodontid fishes, ranging from the Red Sea south to Western Cape, South Africa, east MATERIAL to the western Pacific from New South Wales to Specimens for the present study are from the southern Japan, in Oceania to the Hawaiian Bishop Museum, Honolulu (BPBM), Muséum Islands and islands of French Polynesia and in the National d’Histoire Naturelle, Paris (MNHN), eastern Pacific from Panama to the Gulf of Cali- Universität Humboldt, Museum für Naturkunde, fornia. It is a large species, attaining a total length Berlin (ZMB) and Uppsala Universitet Zoologiska of at least 48 cm. Museet, Uppsala. The similar Arothron reticularis (Bloch & Schnei- We asked representatives of the Senckenberg der, 1801), also a large Indo-Pacific species with a Museum in Frankfurt and the U. S. National type locality of India and ranging east to Fiji Museum of Natural History in Washington, D.C., (Herre 1936), is much less common (for example, to see if their collections held any adult specimens the Bishop Museum has 38 lots of A. hispidus, but of Arothron hispidus with a linear or reticular color only five of A. reticularis). It was not reported from pattern. None were found. the Red Sea until 1956, when Roux-Estève identi- Figures 1-8 are specimen photographs of A. fied a specimen, 197 mm in total length, from the hispidus, in order west to east, from the Red Sea to island of Abulat (Abu Latt) in the Farasan Islands, the Line Islands in the Pacific. Figures 9-17 are southern Red Sea, as Tetraodon reticularis. Her underwater photographs of A. hispidus in the Red record was included in three checklists of Red Sea Sea, and Figures 18-22 are photographs of A. retic- fishes (Dor 1984; Goren & Dor 1994; Golani & ularis, one of which is a juvenile specimen in the Bogorodsky 2010). We cannot find a record of Fisheries Research Laboratory of Mie University, Arothron reticularis from the Indian Ocean west of Japan (FRLM). the southern tip of India (Smith & Heemstra 1986; Fricke 1999; Manilo & Bogorodsky 2003). Arothron hispidus (Linnaeus, 1758) The westernmost locality appears to be Sri Lanka Figures 1-17 (Matsuura in Kimura et al. 2009). Also, A. reticu- laris is often found in estuaries and the young of Tetraodon hispidus Linnaeus, 1758: 333 (type local- Arothron reticularis are restricted to mangrove areas ity, India). and the lower reaches of streams (Veeruraj et al. Tetraodon perspicillaris Rüppell, 1829: 63 (type 2011). The Red Sea coast lacks permanent streams locality, Red Sea). and well-developed estuaries. Roux-Estève’s Red Tetraodon semistriatus Rüppell, 1837: 58, pl. 16, Sea specimen (MNHN 52-252) was examined and fig. 3 (type locality, Massawa, Eritrea). photographed by the third author at the Muséum Tetrodon laterna Richardson, 1845: 124, pl. 61, fig. National d’Histoire Naturelle in Paris (Fig. 1) and 2 (type locality, Canton, China). reidentified as A. hispidus. Tetrodon pusillus Klunzinger, 1871: 645 (type Roux-Estève is not alone in misidentifying speci- locality, El Quseir, Red Sea). mens of Arothron hispidus as A. reticularis. Some Arothron hispidus Matsuura in Masuda et al. 1984: Bishop Museum specimens were first mislabeled as 364, pl. 331 H (Indo-Pacific). A. reticularis, as well as some of the first author’s photographs, and only three of the seven illustra- Diagnosis: Dorsal fin rays 10 or 11; anal fin rays tions currently identified as A. reticularis in FishBase 9-11; pectoral rays 17-19; small spinules on head are correctly identified as this species. Shen et al. and body except for snout, posterior caudal pedun- aqua vol. 18 no. 1 - 15 January 2012 42 John E. Randall, Sergey V. Bogorodsky and Jean Michel Rose

Fig. 1. Arothron hispidus, MNHN 52-252, 160 mm SL, Abu Latt, Farasan Islands. Photo by J. M. Rose.

Fig. 2. Arothron hispidus, BPBM 18112, 240 mm SL, Gulf of Aqaba. Photo by J. E. Randall.

Fig. 3. Arothron hispidus, 333 mm SL, Su dan, specimen lost. Photo by J. E. Randall.

43 aqua vol. 18 no. 1 - 15 January 2012 Color variation of the puffer Arothron hispidus (Linnaeus) and comparison with A. reticularis (Bloch & Schneider)

Fig. 4. Arothron hispidus, BPBM 20139, 80 mm SL, Mauritius. Photo by J. E. Randall.

Fig. 5. Arothron hispidus, BPBM 27686, 210 mm SL, SW India. Photo by J. E. Randall.

Fig. 6. Arothron hispidus, FRLM 30622, 50 mm SL, Libong Island, SW Thailand. Photo by S. Kimura. aqua vol. 18 no. 1 - 15 January 2012 44 John E. Randall, Sergey V. Bogorodsky and Jean Michel Rose cle and base of fins; snout length 4.6-6.1 in stan- base; subadults with white spots often forming cir- dard length (SL); nasal organ with two fleshy flaps cle around eye and adults may have several com- bifurcating from common base; bony ridge above plete white rings (Fig. 11 and Lieske & Myers eye, interorbital space concave; bony interorbital 2004: 220, lower fig.). Subadults (Fig. 4) and juve- width 4.8-7.0 in SL; anal fin origin below or pos- niles (Fig. 6) sparsely spotted with white. With terior to rear base of dorsal fin; caudal fin rounded, growth, white spots become more numerous. Some posterior margin irregular, 2.6-3.8 in SL; color individuals with white lines, as shown in Figures extremely variable, most often olivaceous with 10 and 12, while others (Figs 11,13,14) remaining small white spots, white ventrally, with dark body white-spotted, except for rings around eyes and gill color extending ventrally into the white ventral openings; others developing reticular pattern (Figs part of head and abdomen appearing as bars (three 15 and 16) or a mainly linear pattern (Fig. 17). on head, three on abdomen); base of pectoral fin Base of dorsal fin usually with few white spots; anal within vertically oval black spot containing a few fin generally none. Caudal fin varying from having white or yellow curved lines and/or small spots, only a few white spots on base to being almost often with one or more white lines encircling fin completely white-spotted; some specimens with

Fig. 7. Arothron hispidus, BPBM 14801, 226 mm SL, Lord Howe Island. Photo by J. E. Randall.

Fig. 8. Arothron hispidus, BPBM 33888, 243 mm SL, Kiritimati, Line Islands. Photo by J. E. Randall.

45 aqua vol. 18 no. 1 - 15 January 2012 Color variation of the puffer Arothron hispidus (Linnaeus) and comparison with A. reticularis (Bloch & Schneider)

Fig. 9. Arothron hispidus, Gulf of Aqaba. Photo by J. E. Randall.

Fig. 10. Arothron hispidus, Marsa Alam, Egypt. Photo by S. V. Bogorodsky. aqua vol. 18 no. 1 - 15 January 2012 46 John E. Randall, Sergey V. Bogorodsky and Jean Michel Rose

Fig. 11. Arothron hispidus, Ras Ghamilla, Egypt. Photo by S. V. Bogorodsky.

Fig. 12. Arothron hispidus, Sudan. Photo by J. E. Randall.

Fig. 13. Arothron hispidus, El Quseir, Egypt. Photo by S. V. Bogorodsky.

47 aqua vol. 18 no. 1 - 15 January 2012 Color variation of the puffer Arothron hispidus (Linnaeus) and comparison with A. reticularis (Bloch & Schneider) caudal fins having mixed spots and lines; still oth- reticularis, is the distribution of the small dermal ers with linear or reticular pattern. Largest speci- spinules on the body. They cover the entire body in men reported, 48 cm TL. A. reticularis, except for the region around the Comparison with Arothron reticularis: Because mouth and the base of the fins, whereas they are puffers are so variable in form, it is difficult to find absent on the snout and at least the posterior half morphological characters to distinguish some of the caudal peduncle of A. hispidus (see Fig. 4). species from close relatives. We can confirm that Wheeler (1991: 192, Fig. 29) provided a photo- the most important difference, as reported by Mat- graph of Linnaeus’ holotype of Tetraodon hispidus, suura (1999), in separating A. hispidus from A. ZMUU 102, 93 mm SL, at the University of Upp-

Fig. 14. Arothron hispidus, Sharm el Sheikh, Egypt. Photo by S. V. Bogorodsky.

Fig. 15. Arothron hispidus, Tiran Island, Egypt. Photo by J. L. Rose. aqua vol. 18 no. 1 - 15 January 2012 48 John E. Randall, Sergey V. Bogorodsky and Jean Michel Rose

Fig. 16. Arothron hispidus, El Quseir, Egypt. Photo by S. V. Bogorodsky.

Fig. 17. Arothron hispidus, N Obhur, Saudi Arabia. Photo by R. Field.

49 aqua vol. 18 no. 1 - 15 January 2012 Color variation of the puffer Arothron hispidus (Linnaeus) and comparison with A. reticularis (Bloch & Schneider)

Fig. 18. Arothron reticularis, syntype, ZMB 4259, 185 mm SL, Tranquebar, India. Photo by P. Bartsch.

Fig. 19. Arothron reticularis, BPBM 28931, 355 mm SL, Lizard Island, Great Barrier Reef. Photo by J. E. Randall.

Fig. 20. Arothron reticularis, BPBM 7431, 153 mm SL, Babelthuap, Palau. Photo by J. E. Randall. aqua vol. 18 no. 1 - 15 January 2012 50 John E. Randall, Sergey V. Bogorodsky and Jean Michel Rose

Fig. 21. Arothron reticularis, Baie de Prony, New Caledonia. Photo by R. F. Myers.

Fig. 22. Arothron reticularis, FRLM 30626, 54 mm SL, Libong Island, SW Thailand. Photo by S. Kimura.

51 aqua vol. 18 no. 1 - 15 January 2012 Color variation of the puffer Arothron hispidus (Linnaeus) and comparison with A. reticularis (Bloch & Schneider) sala in Sweden. The spinules are clearly absent tropods, tunicates, sponges, corals, zoanthid from the snout and all of the caudal peduncle. One anemones, crabs, tube worms, sea urchins, brittle of two syntypes of Tetrodon reticularis Bloch & stars, starfishes (including Acanthaster), hermit Schneider at the Museum für Naturkunde der crabs, hydroids and detritus. Humboldt-Universität, Berlin is lost (Paepke The different English common names reflect the 1999). Figure 18 is a photograph of the remaining variation in color pattern: Whitespotted Puffer, syntype, ZMB 4259, 185 mm SL, from Tranque- Stripebelly Puffer, Spiny Balloonfish, and Stars- bar, India. Spinules can be seen on the snout before and-Stripes Puffer. the eye and over the caudal peduncle. Matsuura One Hawaiian name, Makimaki, may be in refer- (1999) also reported that A. reticularis has a wider ence to its toxicity (Titcomb 1972). This species is interorbital space than A. hispidus. one of the most virulently poisonous of tetraodon- Like A. hispidus, A. reticularis is variable in col- tid fishes (Halstead 1967). Yamada in Nakabo oration. It can be distinguished in color by having (2002) reported the skin, testes, liver and ovaries continuous dark brown bands that curve from toxic, adding that there are conflicting reports on below the eye across the abdomen and there are no the toxicity of the flesh. ventral extensions of the dark body color into the Rüppell (1829) described Tetraodon perspicillaris pale ventral part of the head and body. Also, for from the Red Sea, noting the white rings around any given size, the white spots posteriorly on the the orbit and branchial opening. His species name body and caudal fin are larger in A. reticularis and is derived from the Latin perspicillum meaning they more uniformly cover the caudal fin, whereas spectacles, in obvious reference to the rings around in A. hispidus they may be missing from the poste- the eyes. Rüppell (1837: 58, pl. 16, fig. 3) rior part of the fin, if not the entire fin. We present described a second species as Tetraodon semistriatus. color illustrations of an adult of A. reticularis His illustration is the same color form as shown (BPBM 28931, 355 mm SL) from the Great Bar- here by Figure 12 (total length estimated as 32 cm rier Reef (Fig. 19); a subadult (BPBM 7431, 153 when the photo was taken). The second Rüppell mm SL) collected in fresh water from the Mizucho name (semistriatus) is in reference to the body River, Babelthuap, Palau (Fig. 20), a first record for being partly lined. Klausewitz (1960) and Tor- Micronesia; an underwater photograph of an adult tonese (1968) treated the Red Sea population as a taken by Robert F. Myers at Baie de Prony, New subspecies, Arothron hispidus perspicillaris. The Red Caledonia (Fig. 21), a first record for the island; Sea population of Arothron hispidus is more densely and a juvenile from the south-west coast of Thai- white-spotted at a given size (Figs 2, 3, 10-13) than land provided by Seishi Kimura (Fig. 22). that of the Indian Ocean (Figs 4, 5; Smith in Smith Arothron hispidus and A. reticularis are both well & Heemstra 1986: fig. 263; Debelius 1993: 308). illustrated in color in Masuda et al. (1984: pl. As noted by Matsuura in Masuda et al. (1984: 331), Masuda & Kobayashi (1994: 430), and Allen 364, pl. 331, fig. H), Matsuura (1999: 130), and et al. (2003: 441). Figure 7 here, the Pacific Ocean population of Remarks: Arothron hispidus can be found in a Arothron hispidus differs in color from that of the variety of habitats from coral or rocky reefs to open Indian Ocean in having narrow wavy dark stripes sand or mud substrata, seagrass beds, algal flats, or ventrally on the body and fewer white spots. Figure estuaries, from depths of 1-99 m (Struhsaker 8 from Kiritimati in the Line Islands, however, is 1973). It is more often seen in sheltered than an exception in lacking the narrow dark stripes exposed waters, and more often on soft than hard ventrally on the body and in having numerous substrata. It may at times be seen at rest on sand, white spots (as well as bright yellow pectoral fins). algal turf, or seagrass. We have no specimens of large individuals of Bremert & Ormond (1981) reported Arothron Arothron hispidus from the Red Sea that exhibit hispidus feeding in the Red Sea on coral, tunicates, such variable color pattern and express here the sea urchins and brittle stars, adding that it is a sig- need for such specimens. Perhaps some of the vari- nificant predator of the Crown-of-Thorns Starfish ation is a result of sexual dichromatism. Molecular (Acanthaster). They described its mode of feeding study would be most welcome. It would also be on this venomous asteroid in detail. Randall useful in deciding if taxonomic recognition is war- (1985) wrote of the extremely varied diet: algae ranted for the apparent different populations (including Jania and Halimeda), bivalves, gas- within the Indo-Pacific region. aqua vol. 18 no. 1 - 15 January 2012 52 John E. Randall, Sergey V. Bogorodsky and Jean Michel Rose

ACKNOWLEDGEMENTS Sciences and Humanities, Jerusalem and the Interuniver- We thank Dr. Peter Bartsch of the Museum für sity Institute for Marine Sciences, Eilat, 120 pp. HALSTEAD, B. W. 1967. Poisonous and Venomous Marine Naturkunde der Humboldt Universität zu Berlin Animals of the World. Vol. 2 - Vertebrates, United States for his photograph of the existing syntype of Government Printing Office, Washington, D.C., xxxi + Arothron reticularis, Dr. Seishi Kimura of the Fish- 1069 pp. eries Research Laboratory of Mie University for HERRE, A. W. 1936. Fishes of the Crane Pacific Expedi- providing his photographs of juvenile specimens of tion. Zoological Series, Field Museum of Natural History Arothron hispidus and A. reticularis from the 21: 1-472. Andaman Sea, Jean Louis Rose and Richard Field KIMURA, S., SATAPOOMINM, U. & MATSUURA, K. 2009. for their underwater photographs of A. hispidus Fishes of Andaman Sea. National Museum of Nature and Science, Tokyo, vi + 346 pp. taken in the Red Sea, and Robert F. Myers for his KLAUSEWITZ, W. 1960. Systematisch-evolutive Unter- underwater photograph of A. reticularis from New suchungen uber die Abstammung einiger Fische des Caledonia. The most useful reference for this study Roten Meeres. Verhandlungen der Deutschen Zoologischen was the review of the genus Arothron by Keiichi Gesellschaft Münster 1059. Zoologischer Anzeige (Supple- Matsuura (1999), and we thank him for additional ment) 23: 175-182. information. We are grateful to Dr. Tilman Alper- KLUNZINGER, C. B. 1871. Synopsis der Fische des Rothen mann of the Senckenberg Museum, and Dianne Meeres. II. Theil. Verhandlungen der K.-K. zoologisch- Pitassy and Dr. Jeffrey T. Williams of the U. S. botanischen Gesellschaft in Wien v. 21: 441-688. LIESKE, E. & MYERS, R. F. 2004. Coral Reef Guide Red Sea. National Museum of Natural History for searching Harper Collins Publishers Ltd., London, 384 pp. for adult specimens of Arothon hispidus for us. The LINNAEUS, C. 1758. Systema Naturae, ed. X, tom. I, ii + second author acknowledges with gratitude the 824 pp. support for his fieldwork in the Red Sea from the MANILO, L. G. & BOGORODSKY, S. V. 2003. Taxonomic Faculty of Marine Science of King Abdulaziz Uni- composition, diversity and distribution of coastal fishes versity, Jeddah, Saudi Arabia. The third author of the Arabian Sea. Journal of Ichthyology 43(1): 75-149. adds special thanks to Romain Causse, collection MASUDA, H., AMAOKA, K., ARAGA, C., UYENO, T. & manager, Ichthyologie, Muséum National d’His- YOSHINO, T. (eds.) 1984. The Fishes of the Japanese Arch- ipelago. Vol. 1 (text) and vol. 2 (plates), Tokai University toire Naturelle in Paris, for his assistance. Helen K. Press, Tokyo, xxii + 437 pp. Larson, Robert F. Myers and Fareed Krupp MASUDA, H. & KOBAYASHI, Y. 1994. Grand Atlas of Fish reviewed the manuscript and provided very helpful Life Modes. Tokai University Press, Tokyo. 465 pp. (in comments. Japanese). MATSUURA, K. 1999. Taxonomic review of the puffers of the genus Arothron (Tetraodontifomes: Tetraodontidae) REFERENCES with a key to genera of the Indo-West Pacific puffers. ALLEN, G., STEENE, R., HUMANN, P. & DELOACH, N. Ninth Joint Seminar on Marine and Fisheries Sciences, 2003. Reef Fish Identification - Tropical Pacific. New Bali. pp. 125-140. World Publications, Jacksonville, Florida, 457 pp. NAKABO, T. 2002. Fishes of Japan with Pictorial Keys to the BLOCH, M. E. & SCHNEIDER, J. G. 1801. Systema Ichthy- Species. Tokai University Press, Tokyo, English edition, ologiae iconibus cx illustratum. Sanderiano Commissum, vol. 2, vii + 867-1749 pp. Berlin. PAEPKE, H.-J. 1999. Bloch’s Fish Collection in the Museum BREMERT, G. & ORMOND, R. 1981. Red Sea Coral Reefs. für Naturkunde der Humboldt Universität zu Berlin. Kegan Paul International, London and Boston, 192 pp. A.R.G. Gantner Verlag, Liechtenstein, 216 pp. DEBELIUS, H. 1993. Indian Ocean Tropical Fish Guide. RANDALL, J. E. 1985. Guide to Hawaiian Reef Fishes. Harro - Aquaprint, Neu-Isenburg, Germany, 321 pp. wood Books, Newtown Square, Pennsylvania, 74 pp. DOR, M. 1984. CLOFRES: Checklist of the Fishes of the Red RICHARDSON, J. 1845, Ichthyology.—Part 3. In R. B. Sea. The Israel Academy of Sciences and Humanities, Hinds (ed.) The Zoology of the Voyage of H. M. S. Sulphur, Jerusalem. 437 pp. under the Command of Captain Sir Edward Belcher,… FRICKE, R. 1999. Fishes of the Mascarene Islands (Réunion, during the years 1836-42, no. 10. Smith, Elder & Co., Mauritius, Rodriguez). Koeltz Scientific Books, Koenig- London, 51-150 pp. stein, Germany, viii + 759 pp. ROUX-ESTÈVE, R. 1956. Resultats scientifiques des cam- GOLANI, D. & BOGORODSKY, S. V. 2010. The fishes of the pagnes de la “Calypso”. X. Poissons. Annales de l’Institut Red Sea - Reappraisal and updated checklist. Zootaxa Océanographique, Monaco 32, 61-115. 2463: 1-135. RÜPPELL, E. 1828-1829. Atlas zu der Reise im nördlichen GOREN, M. & DOR, M. 1994. An Updated Checklist of the Afrika. Zoologie. Fische des Rothen Meeres. Heinr. Ludw. Fishes of the Red Sea. CLOFRES II. The Israel Academy of Brönner, Frankfurt am Main, 141 pp.

53 aqua vol. 18 no. 1 - 15 January 2012 Color variation of the puffer Arothron hispidus (Linnaeus) and comparison with A. reticularis (Bloch & Schneider)

RÜPPELL, E. 1835-1838. Neue Wirbelthiere zu der Fauna TITCOMB, M. 1972. Native Use of Fish in Hawaii. The Uni- von Abyssinien gehörig. Fische des Rothen Meeres. Frank- versity Press of Hawaii, Honolulu. vii + 175 pp. furt-am-Main. 1-148 [1835: 1-28; 1836: 29-52, 1837: TORTONESE, E. 1968. Fishes from Eilat (Red Sea). Bulletin 53-80; 1838: 1-148]. of the Sea Fisheries Research Station of Haifa Israel 51: 6-30. SHEN, S.-C. (ed.) 1993. Fishes of Taiwan. Department of VEERURAJ, A., ARUMUGAM, M., AJITHKUMAR, T. & BAL- Zoology, National Taiwan University,Taipei, xx + 960 pp. SUBRAMIAN, T. 2011. Distribution of Tetraodontifomes (in Chinese). (Family: Tetraodontidae) along the Paragipettai Coast, SMITH, M. M. & HEEMSTRA, P. C. (eds) 1986. Smiths’ Sea southeast coast of India. Zootaxa 3015: 1-12. Fishes. Macmillan South Africa, Johannesburg, xx + 1047 WHEELER, A. 1991. The Linnaean fish collection in the pp. Zoological Museum of the University of Uppsala. Zoo- STRUHSAKER, P. 1973. A contribution to the systematics and logical Journal of the Linnaean Society 103: 145-195. ecology of Hawaiian bathyal fishes. PhD thesis, University of Hawaii. xv + 482 pp.

aqua vol. 18 no. 1 - 15 January 2012 54 aqua, International Journal of Ichthyology

The first record of Hippocampus denise (Syngnathidae) from Australia

Ralph Foster1, Thomas C. L. Bridge2 and Pim Bongaerts3

1) Ichthyology Section, South Australian Museum, North Terrace, Adelaide, SA 5000, Australia. E-mail: [email protected] 2) School of Earth and Environmental Sciences, James Cook University, Townsville, QLD 4811, Australia. E-mail: [email protected] 3) School of Biological Sciences, University of Queensland, St Lucia, QLD 4072, Australia. E-mail: [email protected]

Received: 18 March 2011 – Accepted: 11 July 2011

Abstract explique que cette espèce a échappé à toute détection dans A specimen of Hippocampus denise (Syngnathidae) la région est due à la connaissance trop partielle des récifs recently collected from the outer Great Barrier Reef, mésophotiques australiens et nous en abordons brièvement Queensland, constitutes the first record of this species les conséquences pour ce qui concerne le statut de la con- from Australian waters. Counts and proportional measure- servation de l’espèce. ments confirm the identity of the specimen. It was taken by ROV at a depth greater than previously reported for the Sommario species and from a new host species of octocoral. We pos- Un esemplare di Hippocampus denise (Syngnathidae) re- tulate that one reason for the species having previously cen temente raccolto lungo il margine esterno della Grande evaded detection in the region is due to the paucity of Barriera Australiana, Queensland, rappresenta la prima se- knowledge of Australian mesophotic reef systems and gnalazione di questa specie in acque australiane. Le misu - briefly discuss the implications of this on assessing the razioni meristiche confermano l’identificazione dell’esem- species’ conservation status. plare. Questo è stato raccolto con l’ausilio di un ROV ad una profondità più elevata di quella precedentemente regi - Zusammenfassung strata per questa specie e su una diversa specie ospite di otto- Kürzlich konnte ein Exemplar von Hippocampus denise corallo. Si ipotizza che una ragione del mancato rilevamento (Syngnathidae) im äußeren Bereich des Großen Barri- di questa specie nella regione sia dovuto alla scarsa ereriffs, Queensland, gefangen werden; damit ist der Erst- conoscenza dei sistemi mesofotici della barriera australiana e nachweis dieser Art in australischen Gewässern gelungen. si discutono brevemente le implicazioni che ciò può com- Nach Kennzahlen und proportionalen Messungen konnte portare nella determinazione dello stato di conservazione die Art eindeutig bestimmt werden. Das Exemplar war delle specie. durch ein ferngesteuertes Fahrzeug (ROV) in einer größeren Tiefe gesammelt worden als nach bisherigen INTRODUCTION Berichten und über einer neuen Wirtsart der Octoko- rallen. Ein Grund für den bisher fehlenden Nachweis dort One of the smallest known species of seahorse, sind unserer Auffassung nach die geringen Kenntnisse über Hippocampus denise Lourie & Randall, 2003, has die mesophotischen australischen Riffsysteme; die Fol- not been recorded from Australia despite its very gerungen im Blick auf den Schutzstatus der Art werden widespread occurrence in adjacent regions of the kurz diskutiert. Indo-West Pacific that make up the “Coral Trian- gle”. This gorgonian octocoral-dwelling species is Résumé recorded to range from Malaysia and throughout Un spécimen d’Hippocampus denise (Syngnathidae) Indonesia to Palau, Micronesia and south-east to récemment collecté au large de la Grande Barrière de the Solomon Islands and Vanuatu (Lourie & Ran- Corail, Queensland, constitue la première mention de cette espèce pour les eaux australiennes. Les données dall 2003). Lourie & Kuiter (2008) also report it méristiques et morphométriques confirment l’identité de from New Ireland Province, Papua New Guinea l’espèce. Elle a été capturée par ROV à une profondeur (PNG) and photographs published on the web plus importante que signalée jusqu’ici et sur une nouvelle indicate that it also occurs in West New Britain espèce-hôte d’octocorail. Nous postulons que la raison qui and Milne Bay Provinces, PNG (Banks 2006 –

55 aqua vol. 18 no. 1 - 15 January 2012 The first record of Hippocampus denise (Syngnathidae) from Australia reported as H. bargibanti Whitley). We report the 31.7); TaL 57.7% in SL (47.4-57.2); TD9 10.2% first finding of Hippocampus denise in Australian in SL (4.1-15.5); DL 10.2% in SL (6.4-10.1). waters, from a single specimen on an octocoral col- In appearance the specimen SAMA F13806 lected by a remotely operated vehicle (ROV) used closely resembles the description and published in a study of mesophotic (30-150 m depth) coral photographs of Hippocampus denise (Lourie & communities off the North Queensland coast Randall 2003). All counts and proportional mea- (Bongaerts et al. 2011). It is deposited in the South surements are consistent with those reported for Australian Museum, Adelaide (SAMA). the species by Lourie & Randall (2003) and Lourie & Kuiter (2008) with the departures of TaL and MATERIAL AND METHODS DL from the cited ranges statistically insignificant. Material examined – Hippocampus denise, SAMA A similar species, H. bargibanti, is known to occur F13806, 13.7 mm SL, approximately 240 km on the Great Barrier Reef (GBR) (Paxton et al. ENE Cairns, Qld, NW corner of Holmes Reef 2006) but significant proportional measurements East, 16°.421’ S, 147°.987’ E, 25 October 2010, (Lourie & Randall 2003: Table I) along with tail on Villogorgia sp. taken by ROV from a depth of ~100 metres, collected by Tom Bridge and Dave Whillas (AUV Coral Sea Exploratory Expedition); fixed and preserved in 70% ethanol. To verify the specimen’s identity, counts and mea- surements were taken following Lourie & Randall (2003). Counts of trunk rings (TrR), tail rings (TaR) and pectoral, dorsal and anal fin rays (PF, DF and AF, respectively) were made using a micro- scope and from a microtomagraph (CT scan) of the specimen as per Foster & Gomon (2010). Mea- surements to the nearest 0.1 mm were made using a microscope with an eyepiece graticule and from an enlarged photograph incorporating a scale bar. Measurements were made of head length (HL) trunk length (TrL) tail length (TaL), snout length (SnL), orbital diameter (OD), post orbital length (PO), crown height (CH) trunk depth at TrR 9 (TD9), dorsal fin base length (DL). Standard length (SL) = HL+TrL+TaL.

RESULTS AND DISCUSSION Description - Colour in life, dark reddish-orange with faint banding on the tail; sex uncertain but body proportions and comparison of the CT scan with X-rays of sexed individuals (Lourie & Randall 2003) suggest it is a non-breeding male. Counts – TrR 12; TaR 29; PF 11; DF 14; AF 4. Principal measurements (mm) – HL 2.2; TrL 3.6; TaL 7.9. Proportional measurements (as percent- ages of SL, HL or SnL, as indicated, with range from 14 specimens of Hippocampus denise exam- ined by Lourie & Kuiter (2008) shown in paren- theses) – HL 16.1% in SL (16.0-23.1); HD 53.6% in HL (41.1-55.7); SnL 37.2% in HL (27.1-38.7); SnD 76.4% in SnL (62.7-81.2); OD 23.5% in HL Fig. 1. Hippocampus denise, SAMA F13806, from Holmes (18.8-23.5); PO 40.8% in HL (39.1-45.2); CH Reef East, Queensland photographed soon after collection. 45.4% in HL (35.2-51.6); TrL 26.3% in SL (23.1- Photo by E. Roberts. aqua vol. 18 no. 1 - 15 January 2012 56 Ralph Foster, Thomas C. L. Bridge and Pim Bongaerts ring count, a non-bulbous snout and an absence of Museum for CT scanning the specimen, Chris tubercles above the eyes specifically exclude this Jones for discovering the specimen on the octoco- species. ral, David Whillas for the ROV operations, Ed As well as being the first Australian record of Hip- Roberts for photographing the specimen, and the pocampus denise, the specimen, SAMA F13806, is crew of Eye to Eye Marine Encounters for their notable for the depth at which it was found (100 logistical support. Thanks also to the anonymous m) - the deepest yet reported for the species - and reviewers for their very helpful comments on the for the host species, Villogorgia sp. Previously manuscript. We acknowledge Australian Geo- reported hosts are Annella, Muricella and Echino- graphic, C&R Consulting, the National Science gorgia, at depths ranging from 13-90 m (Lourie & Foundation (ATM-0941760) and the Pacific Blue Randall 2003). Gorgonian octocorals occur on Foundation for funding. mesophotic reefs throughout northern Australian seas, including a vast area of the GBR outer shelf to 150 m depth (Bridge et al. 2011). That H. denise REFERENCES BANKS, I. 2006. Diving the Gold Coast with Ian Banks has not previously been recorded seems surprising http://www.divingthegoldcoast.com.au/index.asp?PageI and suggests that it may be rare in Australian D=animal&CritterID=2109 waters. However, this species of pygmy seahorse, http://www.divingthegoldcoast.com.au/index.asp?PageI along with others that may occur in the region, is D=animal&CritterID=2417 so small and cryptic that it is extremely difficult to http://www.divingthegoldcoast.com.au/index.asp?PageI find, the difficulty compounded by its distribution D=animal&CritterID=2430 well into the mesophotic zone beyond the range of http://www.divingthegoldcoast.com.au/index.asp?PageI scuba divers. Mesophotic communities in Australia D=animal&CritterID=5064 have received little attention compared to their Accessed 2 Dec 2010. BONGAERTS, P., BRIDGE, T. C. L., KLINE, D. I., MUIR, shallow-water counterparts due to their relative P. R., WALLACE, C. C., BEAMAN, R. J. & HOEGH-GUL- inaccessibility and only recently have technological BERG, O. 2011. Mesophotic coral ecosystems on the walls advances, such as the use of ROVs, opened up the of Coral Sea atolls. Coral Reefs 30: 335. zone for exploration. Well-known Coral Triangle BRIDGE, T. C. L., DONE, T. J., BEAMAN, R. J., FRIED- octocoral species have also recently been found for MAN, A., WILLIAMS, S. B., PIZARRO, O. & WEBSTER, the first time in Australian waters during surveys of J. M. 2010. Topography, substratum and benthic macro- the GBR outer shelf (Bridge, unpublished data) faunal relationships on a tropical mesophotic shelf mar- gin, central Great Barrier Reef, Australia. Coral Reefs 30: and it is probable that more species of pygmy sea- 143-153. horses occur in northern Australian waters but no FOSTER, R. & GOMON, M. F. 2010. A new seahorse focused searches have yet been made. Unfortu- (Teleostei: Syngnathidae: Hippocampus) from south- nately, research is hampered in Australia by restric- western Australia. Zootaxa 2613: 61-68. tive scientific diving regulations on, for example, LOURIE, S. A. & KUITER, R. 2008. Three new pygmy sea- the use of rebreather units which are being used horse species from Indonesia (Teleostei: Syngnathidae: elsewhere to survey the mesophotic zone (e.g. Pyle Hippocampus) Zootaxa 1963: 54-68. et al. 2008). LOURIE, S. A. & RANDALL, J. E. 2003. A new pygmy sea- horse, Hippocampus denise (Teleostei: Syngnathidae) Pygmy seahorses are tiny, few more so than H. from the Indo-Pacific. Zoological Studies 42 (2): 284-291. denise, and are likely to be missed unless specifically PAXTON, J. R., GATES, J. E., HOESE, D. F. & BRADY, D. J. searched for in suitable communities. Hippocampus 2006. Syngnathidae In: HOESE, D. F., BRAY, D. J., PAX- denise is listed as “Data Deficient” on the IUCN TON, J. R. & ALLEN, G. R. Zoological Catalogue of Red List of Threatened Species (Project Seahorse Australia Volume 35 Fishes. (eds P. L. Beesley & A. Wells) 2003) and the conservation action calls for further Part 2 pp 810-846. ABRS & CSIRO Publishing: research on the species biology, ecology, habitat, Australia. abundance, and distribution. Targeted surveys are PROJECT SEAHORSE 2003. Hippocampus denise. In: IUCN 2011. IUCN Red List of Threatened Species. Version necessary to gather this information before an 2011.1. . Accessed on 17 June 2011 assessment of the conservation status of the species PYLE, R. L., EARLE, J. L. & GREENE, B. D. 2008. Five new in Australia can be made. species of the damselfish genus Chromis (Perciformes: Labroidei: Pomacentridae) from deep coral reefs in the ACKNOWLEDGEMENTS tropical western Pacific. Zootaxa 1: 3-31 Thanks go to Peter Blias of the South Australian

57 aqua vol. 18 no. 1 - 15 January 2012 Teodor T. Nalbant – 1933-2011

“The foundation of my love of science is my water and marine fishes of Romania, then on love of beauty” (Cyril Crossland) was Teodor the ichthyofauna from other geographical Nalbant’s motto for his scientific career. areas – his first paper appeared in 1956. He “These are simple words which are better covered a very large spectrum of fields: sys- understood only by those who dedicated years of tematics, morphology, speciation, ecology, their lives to the biological field” he said during ethology, zoogeography, paleontology, and the celebration of his 70th anniversary, while other aspects of an applied nature. he had reached 50 years of activity within the But one of Nalbant’s main concern, was the scientific community. study of the ichthyofauna based on the mate- Nalbant was born on 18th of December rial collected by Romanian and foreign expe- 1933, on the shore of the Black Sea, at Con- ditions. Within this context his studies stants, Romania. Moving with his family to resulted in very important discoveries in Tulcea in 1939, a decisive moment for his marine fishes from the Central and North future profession and life plans began with the meeting of Pacific, Eastern and North-Western tropical Atlantic, the the Danube Delta where I had my first meeting with this Indian Ocean, and freshwater fishes from Central Africa, unforgettable scientist. We did a field trip together in 2008 Afghanistan, the Himalayas, India, Iran, Turkey, and Cuba. to the place where it had all begun for him. Prior we were Also his studies on fishes of the family Chaetodontidae were invited for a seminar in Bucharest’s “Grigore Antipa” of great importance, from material mostly collected during National Museum where the monumental B n rescu – the expeditions of the “Uranie“, “Challenger“, “Albatross“, Nalbant collection with over 10,000 specimens, especially “Galathea“, and “Noona Dan”. of the families Cyprinidae, Cobitidae and Gobiidae, is As a result of his studies, he discovered and described, under the custody of the Museum. We also went together alone or in collaboration a new cobitid subfamily (Vail- to the type locality of the endemic genus Romanichthys val- lanelinae Nalbant, B n rescu, 1997 – Cobitidae), and an sanicola and had a great time together. Two years later he impressive number of taxa whose illustrations point out not came to visit me in Italy and we agreed to do a book only his talent in drawing (sample below) but also in accu- together, about the cobitids and nemacheilids of the world rate observations. His descriptions include 23 new genera and I got started with specimens I collected for him in and subgenera, and 58 species and subspecies including a Turkey and elsewhere in 2009. Unfortunately his health blind fish, Lucifuga simile. Nalbant contributed to many must have been deteriorating, I rarely heard from him. He international publications by writing the chapters on the passed away while I was on field trips in the Amazon on families Cobitidae and Chaetodontidae. He participated in November 29th, 2011. long lasting expeditions as in the North-West Atlantic Nalbant’s family wanted him to follow a medical career, Ocean (in 1967 and 1969) and along the West African but he succeeded in attending the Faculty of Biology from coasts (1969), as well as in Romanian expeditions (one with Clujand in the autumn of 1953, he met Petru Bănărescu me) and one to the coast of Tanzania resulting in the dis- with whom he continuously shared, till Bănărescu’s end covery and the description of numerous new genera and (1921-2009), his love of beauty and fishes, publishing species. together more than 40 papers, especially in the systematic, Nalbant always correlated the outer morphological features phylogeny and zoogeography of the fishes of the families with those of the inner morphology, of ecology, ethology Cyprinidae (Gobioninae) and Cobitidae. In February 1958 and of zoogeography which was and is very important for a Nalbant graduated from the University of Bucharest, and better understanding of species. Something all ichthyolo- then worked as a biologist at the Institute of Hydrotechni- gists should do. The genus Nalbatichtlzys Schultz 1967, as cal Studies and Research (1959- 1961), followed by the well the subgenus Nalbantius Maugé & Bauchot, 1984 Institute of Fisheries (1961-1970), where he became the (later given generic status), Schistura nalbarzti Mirza & senior scientist. Later he was employed by the National B n rescu, 1979, and the cumacean crustacean Cyclaspis nal- Museum of Natural History in Bucharest “Grigore Antipa” hanti Petrescu, 1998, immortalized this amazing scientist (1970-l975), where he was senior curator, in the Institute of and certainly no ichthyologist or loach-lover will ever forget Biological Sciences, Department of Taxonomy and Evolu- this humble and very natural Romanian. Farewell Theo. tion. Nalbant published over 150 papers, first on the fresh- Heiko Bleher

aqua vol. 18 no. 1 - 15 January 2012 58 Helen Larson – retired from aqua

What can I say about the woman who Helen, I look forward to your having probably knows more about the family the time you want and need during Gobiidae than anyone else? What can I your retirement to complete the say about Helen Larson that the ichthy- tremendous task of recording most of ological society and scientists in this the missing species of the largest fish world do not know already? I am sure family on this planet, not only to cata- not much. But there is something I loguing and giving them names, but must say, and that is: THANK YOU also to revising those groups which are VERY MUCH for the fantastic work in limbo. I promise to help as long as I you have done for three consecutives can and in every possible way. I look years for this journal as scientific editor forward to our project with Doug and (2009-2011). THANK YOU for hav- hope it will take off and mature in ing helped so many to publish only 2012. I look forward to continuing to prime peer-reviewed articles in aqua work with you to compile the volumes and in rejecting those which have not of the Gobioidei group, and to having deserved to be published. THANK in a few years something extremely YOU for the advice you gave to authors valuable for today’s and future genera- around the globe, for all the corrections tions. For ichthyologists, taxonomists and scientists, and you have done, and especially on some very long papers. for all people interested in learning more about the amaz- THANK YOU for your tireless and constant communica- ing goby-treasures we have on earth, … thank you. tion with reviewers worldwide. Heiko Bleher

Frank L. Pezold III – new Scientific Editor of aqua

I am also very thankful to Frank L. tion and adaptive radiation of gobies, 4) Pezold III, for taking over as the new sci- cryptic speciation in African freshwater entific editor for aqua, with the begin- sleepers (Eleotridae: Kribia), and 5) con- ning of 2012 and volume 18. servation genetics of California desert Frank has studied at the University of fishes. His research is been supported by New Orleans, B.A. English (1974), M.S. the National Science Foundation in Biological Sciences (1979), and at the (NSF), the Mac Arthur Foundation, University of Texas at Austin, where he World Bank, the USDA Forest Service, received his Ph.D. in Zoology (1984). the Nature Conservancy and Conserva- Today he is Professor of Biology and tion International. And Frank’s recent Dean of the College of Science and Engi- publications have appeared in Zootaxa, neering at Texas A&M University – Cor- Biology of Fishes, Journal of Natural pus Christi (TAMUCC) (since 2006). History, at Science Publisher Inc., in He has also been a Research Associate of aqua, International Journal of Ichthyol- the American Museum of Natural His- ogy, in Fishes of the freshwaters of the tory since 2005. lower Guinea ichthyogeographical Frank’s lab focuses on the diversity, evo- province: Cameroon, Equatorial Guinea, lution and conservation of fishes with an emphasis on gob- Gabon and Congo Brazzaville. G. Teugels, M. Stiassny and ioid fish systematics. He and his students conduct fieldwork C. Hopkins eds., in the Proceedings of the California Acad- in the southern US, California, Mexico, Central America, emy of Sciences and in Copeia. Caribbean Sea, West Africa, Red Sea, and throughout the I look forward to working very closely together with Frank central, western and southern Pacific nations. His current for aqua to grow even more in the years to come, although projects (besides aqua) include: 1) population structure of it is already today one of the prime scientific journal of amphidromous fishes of Micronesia, 2) evolution and sys- ichthyology, thanks to the great help from recent scientific tematics of the gobiid genera Ctenogobius, Oxyurichthys, editors, the late Keith Banister, Walter Ivantsoff, Friedhelm, Stiphodon and Sicydium, 2) cryptobenthic Pacific coral reef Krupp, and Helen Larson. fish community ecology and evolution, 3) ecological specia- Heiko Bleher

59 aqua vol. 18 no. 1 - 15 January 2012 Index of aqua Vol. 17 (1-4) (Index by: 1. Author(s); 2. New Taxa; 3. Biology/Ecology/Biography/Reviews) Author(s): Abitia-Cárdenas,Leonardo A., Moreno-Sánchez, Xchel G., Palacios-Salgado, Deivis S. and Escobar-Sánchez, Ofelia: Feeding habits of the convict surgeonfish Acanthurus triostegus (Teleostei: Acanthuridae) on the Los Frailes reef, Baja California Sur, Mexico. aqua 17 (3): 121-126, 10 July 2011 Allen, Gerald R., Erdmann, Mark V. and Hiloman, Vincent V.: A new species of damselfish (Pomacentrus: Pomacentridae) from Brunei and the Philippines. aqua 17 (1): 35-42, 15 January 2011 Bejarano-Álvarez, Marcela, Galván-Magaña, Felipe and Ochoa-Báez, Rosa Isabel: Reproductive biology of the scalloped hammerhead shark Sphyrna lewini (Chondrichthyes: Sphyrnidae) off south-west Mexico. aqua 17 (1): 11-22, 15 January 2011 Castro, José I.: Resurrection of the name Carcharhinus cerdale, a species different from Carcharhinus porosus. aqua 17 (1): 1-10, 15 January 2011 Clark, Eugenie and Randall, John E.: Cephaloscyllium stevensi: a new species of swellshark (Carcharhiniformes: Scyliorhinidae) from Papua New Guinea. aqua 17 (1): 23-34, 15 January 2011 Hadiaty, Renny K. and Allen, Gerald R.: Glossamia arguni, a new species of freshwater cardinalfish (Apogonidae) from West Papua Province, Indonesia. aqua 17 (3): 173-180, 10 July 2011 Hoese, Douglass F., Winterbottom, Richard and Reader, Sally: Trimma maiandros, a new species of pygmy goby (Gobiidae) from the Indo-west Pacific. aqua 17 (2): 103-110, 26 April 2011 Hoese, Douglass F., Shibukawa, Koichi and Sakaue, Jiro: A redescription of the gobiid fish Cryptocentrus sericus Herre, with clarification of Cryptocentrus leptocephalus and C. melanopus. aqua 17 (3): 163-172, 10 July 2011 Ivantsoff, Walter and Allen, Gerald R.: A new species and genus of a large and unusual freshwater hardyhead, Sashatherina giganteus (Pisces: Atherinidae) from West Papua, Indonesia and a comparison with its closest relatives of the genus Craterocephalus. aqua 17 (1): 43-57, 15 January 2011 Knuckey, James D. S., Ebert, David A. and Burgess, George H.: Etmopterus joungi n. sp., a new species of lanternshark (Squaliformes: Etmopteridae) from Taiwan. aqua 17 (2): 61-72, 26 April 2011 Kullander, Sven O.: A review of Dicrossus foirni and Dicrossus warzeli, two species of cichlid fishes from the Amazon River basin in Brazil (Teleostei: Cichlidae). aqua 17 (2): 73-94, 26 April 2011 Riesch, Rüdiger, Colston, Timothy J., Joachim, Brandon L. and Schlupp, Ingo: Natural history and life history of the Grijalva gambusia Heterophallus milleri Radda, 1987 (Teleostei: Poeciliidae). aqua 17 (2): 95-102, 26 April 2011 Valdesalici, Stefano and Amato, Giuseppe: Nothobranchius oestergaardi (Cyprinodontiformes: Nothobranchiidae), a new annual killifish from Mweru Wantipa Lake drainage basin, northern Zambia. aqua 17 (2): 111-119, 26 April 2011 Winterbottom, Richard: Six new species of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature. aqua 17 (3): 127-162, 10 July 2011

New Taxa: Cephaloscyllium stevensi n. sp. A new species of swellshark (Carcharhiniformes: Scyliorhinidae) from Papua New Guinea. aqua 17 (1): 23-34, 15 January 2011 Etmopterus joungi n. sp. A new species of lanternshark (Squaliformes: Etmopteridae) from Taiwan. aqua 17 (2): 61-72, 26 April 2011 Glossamia arguni n. sp. A new species of freshwater cardinalfish (Apogonidae) from West Papua Province, Indonesia. aqua 17 (3): 173-180, 10 July 2011 Nothobranchius oestergaardi n. sp. (Cyprinodontiformes: Nothobranchiidae), a new annual killifish from Mweru Wantipa Lake drainage basin, northern Zambia. aqua 17 (2): 111-119, 26 April 2011 Pomacentrus cheraphilus n. sp. A new species of damselfish (Pomacentrus: Pomacentridae) from Brunei and the Philippines. aqua 17 (1): 35-42, 15 January 2011 Sashatherina n. gen. A new species and genus of a large and unusual freshwater hardyhead, Sashatherina giganteus (Pisces: Atherinidae) from West Papua, Indonesia and a comparison with its closest relatives of the genus Craterocephalus. aqua 17 (1): 43-57, 15 January 2011 Sashatherina giganteus n. sp. A new species and genus of a large and unusual freshwater hardyhead, Sashatherina giganteus (Pisces: Atherinidae) from West Papua, Indonesia and a comparison with its closest relatives of the genus Craterocephalus. aqua 17 (1): 43-57, 15 January 2011 Trimma cheni n. sp. Six new species of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature. aqua 17 (3): 127-162, 10 July 2011 Trimma erdmanni n. sp. Six new species of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature. aqua 17 (3): 127-162, 10 July 2011 Trimma habrum n. sp. Six new species of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature. aqua 17 (3): 127-162, 10 July 2011 Trimma haimassum n. sp. Six new species of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature. aqua 17 (3): 127-162, 10 July 2011 Trimma maiandros n. sp. A new species of pygmy goby (Gobiidae) from the Indo-west Pacific. aqua 17 (2): 103-110, 26 April 2011 Trimma papayum n. sp. Six new species of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature. aqua 17 (3): 127-162, 10 July 2011 Trimma xanthochrum n. sp. Six new species of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature. aqua 17 (3): 127-162, 10 July 2011

Biology/Ecology/Biography/Reviews: A redescription of the gobiid fish Cryptocentrus sericus Herre, with clarification of Cryptocentrus leptocephalus and C. melanopus. aqua 17 (3): 163-172, 10 July 2011 A review of Dicrossus foirni and Dicrossus warzeli, two species of cichlid fishes from the Amazon River basin in Brazil (Teleostei: Cichlidae). aqua 17 (2): 73-94, 26 April 2011 Book review: The Lady and the Sharks by Eugenie Clarke. aqua 17 (1): 58, 15 January 2011 Book review: The life of an ichthyologist by A. P. Saveliev. aqua 17 (1): 59, 15 January 2011 Feeding habits of the convict surgeonfish Acanthurus triostegus (Teleostei: Acanthuridae) on the Los Frailes reef, Baja California Sur, Mexico. aqua 17 (3): 121-126, 10 July 2011 Natural history and life history of the Grijalva gambusia Heterophallus milleri Radda, 1987 (Teleostei: Poeciliidae). aqua 17 (2): 95-102, 26 April 2011 Reproductive biology of the scalloped hammerhead shark Sphyrna lewini (Chondrichthyes: Sphyrnidae) off south-west Mexico. aqua 17 (1): 11-22, 15 January 2011 Resurrection of the name Carcharhinus cerdale, a species different from Carcharhinus porosus. aqua 17 (1): 1-10, 15 January 2011 aqua vol. 18 no. 1 - 15 January 2012 60 Guidelines for Authors 1. Manuscript preparation: manuscripts must be submit- Examples of correct reference formats: ted in English. In exceptional cases aqua may provide BLABER, S. J. M. 1980. Fish of the Trinity inlet system of translations of manuscripts written in French, German, North Queensland, with notes on the ecology of fish Italian, or Spanish. faunas of tropical Indo-Pacific estuaries. Australian Manuscripts must be word-processed in Microsoft Journal of Marine and Freshwater Research 31:137-46. WORD and submitted in an electronic form. Generic, specific, and sub-specific names must be italicised. All DAY, J. H., BLABER, S. J. M., & WALLACE, J. H. 1981. papers must conform to the International Code of Zoo- Estuarine fishes. In: Estuarine Ecology with Particular logical Nomenclature. Authors are strongly advised to fol- Reference to Southern Africa. (Ed. J.H. Day.): 197- low the format set out in previous publications of aqua. 221. A. A. Balkema, Rotterdam. 2. Title: the title must be short and should precisely iden- DIMMICH, W. W. 1988. Ultrastructure of North Ameri- tify the main topic of the article. Names of genera or can cyprinid maxillary barbels. Copeia 1988 (1): 72- species are followed by the systematic group to which they 79. belong. Author name(s) are given in full beneath the title, TREWAVAS, E. 1983. Tilapiine Fishes of the Genera followed by the complete mailing and e-mail address(es). Sarotherodon, Oreochromis and Danakilia. British 3. Abstract: the abstract should not exceed 250 words and Museum (Natural History), London, 583 pp. draw attention to the principal conclusions. It should not contain any uncommon abbreviations or literature cita- 6. Submission of manuscript and illustrations: The man- tions. The inclusion of abstracts in other languages is uscript and illustrations must be submitted digitally to the optional. Scientific Editor: 4. Subject matter: the text of the manuscript is usually Dr Frank L. Pezold arranged in four main sections: Introduction, Materials College of Science & Engineering and methods (including a key to abbreviations), Results, Texas A&M University – Corpus Christi and Discussion. Other subdivisions may be chosen 6300 Ocean Drive depending on the material presented. Acknowledgements Corpus Christi, TX 78412-5806 should be placed between the text and references. Email: [email protected] Scientific names of genera, species, and subspecies should be followed by the name(s) of author(s) and the year of to whom all subsequent correspondence shall be addressed. publication on first mention. A description of a new taxon Texts, tables, and graphs should be in Microsoft-compati- must contain the following sections: Material, Diagnosis, ble electronic form. Description, and Affinities. Synonyms must be arranged After the paper has been accepted for publication, illustra- in chronological order. Identification keys must be tions as high-resolution TIF files or original photographs dichotomous. (ideally transparencies; otherwise glossy prints, preferably Holotype and paratypes must be clearly identified, the in the size in which they will appear - the type area of aqua institution in which they have been deposited named, and is 158 x 224 mm, one column is 76 mm wide) must be the catalogue numbers given. Private collections are not sent to: acceptable as repositories for holotypes. Aquapress, The Managing Editor DNA sequences must be archived in GenBank, DNA Via G. Falcone 11, Databank of Japan or European Molecular Biology Labora- 27010 Miradolo Terme (Pavia), Italy tory. Voucher specimens associated with DNA sequences E-mail: [email protected] must be deposited in a recognized research collection acces- Authors should retain copies of all materials for reference. sible to the professional community. Accession numbers for sequences and catalog number for vouchers must be Proofs of accepted papers will be sent as PDF files by e- included as a table in an appendix to the final manuscript. mail attachment to the corresponding author. The metric system and SI units must be used. Tempera- 7. Evaluation of manuscripts: manuscripts will be evalu- tures are given in °C. Fractions should not be used. ated by the editors and referees. Papers are accepted on the understanding that they have not and will not be pub- 5. References to literature: the name-year system must be lished elsewhere. used. The list of references should be placed at the end of the paper, alphabetically arranged according to author 8. Reprints: Authors will receive one free copy of the issue name. Only those publications cited in the paper may be in which their paper appears and an e-print in PDF for- included. Titles of journals must be given in full. mat. Additional copies may be ordered from Aquapress. aqua International Journal of Ichthyology Vol. 18 (1), 15 January 2012

Contents:

Fenton Walsh and Hiroyuki Tanaka: Cirrhilabrus nahackyi, a new wrasse (Perciformes; Labridae) from the South Pacific...... 1-8 Gerald R. Allen and Mark V. Erdmann: A new species of Dragonet (Synchiropus: Callionymidae) from Indonesia ...... 9-14 E. Mauricio Hoyos-Padilla, B. Patricia Ceballos-Vázquez and Felipe Galván-Magaña: Reproductive Biology of the Silky Shark Carcharhinus falciformis (Chondrichthyes: Carcharhinidae) off the west coast of Baja California Sur, Mexico ...... 15-24 William D. Anderson, Jr. and Graciela García-Moliner: A new species of Odontanthias Bleeker (Perciformes: Serranidae: Anthiinae) from Mona Passage off Puerto Rico, the first record of the genus from the Atlantic Ocean ...... 25-30 Samaneh Poursaeid & Bahram Falahatkar: Threatened fishes of the world: Stenodus leucichthys leucichthys Güldenstädt, 1772 (Salmonidae) ...... 31-34 John E. McCosker and Gerald R. Allen: Description of a new Snake Eel (Pisces: Ophichthidae: Myrichthys) from the Philippines ...... 35-40 John E. Randall, Sergey V. Bogorodsky and Jean Michel Rose: Color variation of the puffer Arothron hispidus (Linnaeus) and comparison with A. reticularis (Bloch & Schneider) ...... 41-54 Ralph Foster, Thomas C. L. Bridge and Pim Bongaerts: The first record of Hippocampus denise (Syngnathidae) from Australia ...... 55-57 Orbituary: Teodor T. Nalbant (1933-2011) ...... 58 Dr Helen Larson – retired from aqua and Dr Frank L. Pezold – new Scientific Editor of aqua ...... 59 Index of aqua Vol. 17 (1-4) ...... 60

Papers appearing in this journal are indexed in: Zoological Record; BioLIS – Biologische Literatur Information Senckenberg; www.aqua-aquapress.com; www.aquapress-bleher.com; www.Joachim-Frische.com

Cover photo: Comparison of head coloration of Myrichthys paleracio. Photos by G. R. Allen

New genus and new species of Gobiidae in a upcoming issue of aqua. Photo by J. Van Tassell.