Aqua, International Journal of Ichthyology Aqua, International Journal of Ichthyology

aqua International Journal of Ichthyology

Vol. 19 (3), 19 July 2013

Aquapress ISSN 0945-9871 aqua - International Journal of Ichthyology Managing Editor: Scope aqua is an international journal which publishes original Heiko Bleher scientific articles in the fields of systematics, taxonomy, Via G. Falcone 11, bio geography, ethology, ecology, and general biology of 27010 Miradolo Terme (PV), Italy fishes. Papers on freshwater, brackish, and marine fishes Tel. & Fax: +39-0382-754129 will be considered. aqua is fully refereed and aims at pub- E-mail: [email protected] lishing manuscripts within 2-4 months of acceptance. In www.aqua-aquapress.com view of the importance of color patterns in species identi - fication and animal ethology, authors are encouraged to submit color illustrations in addition to descriptions of Scientific Editor: coloration. It is our aim to provide the international scientific community with an efficiently published journal Frank Pezold meeting high scientific and technical standards. College of Science & Engineering Texas A&M University – Corpus Christi Call for papers 6300 Ocean Drive – Corpus Christi, TX 78412-5806 The editors welcome the submission of original manu- Tel. 361-825-2349 scripts which should be sent in digital format to the scien- E-mail: [email protected] tific editor. Full length research papers and short notes will be considered for publication. There are no page charges and color illustrations will be published free of charge. Authors will receive one free copy of the issue in which Editorial Board: their paper is published and an e-print in PDF format. Gerald R. Allen Department of Aquatic Zoology, Subscription Notice Western Australian Museum, Perth, Australia At least one volume (4 issues) of aqua is being published per year, each issue comprising 38-64 pages (incl. cover). Nina G. Bogutskaya The subscription rate (for one volume = 4 issues): Zoological Institute of the Russian Academy of Personal subscription: Euro 75,00 (online edition) - Euro Sciences, St. Petersburg, Russia 100,00 (online edition + print edition incl. priority mail); Institutional subscription: Euro 800,00 (online edition) - Friedhelm Krupp Euro 1000,00 (online edition + print edition incl. priority Curator of Fishes, Senckenberg Research Institute mail). Subscription enquires should be sent to the and Natural History Museum, Frankfurt am Main, publisher at the address given below or by e-mail to: Germany [email protected] - [email protected] Flávio C. T. Lima Special Publication Museu de Zoologia da Universidade de São Paulo Since 2003 Aquapress publishes a series of Special Publi- São Paulo, Brasil cations, which are produced at irregular intervals. All Spe- cial Publications have about 100 or more pages and are Axel Meyer available separately from regular issues of aqua. Enquiries Lehrstuhl für Zoologie und Evolutions biologie, about subscriptions and prices should be sent to the pub- Universität Konstanz, Germany lisher at the address given here above or by e-mail to: [email protected] - [email protected] Paolo Parenti Department of Enviromental Sciences, University of Milano-Bicocca, Milan, Italy Mário de Pinna Museu de Zoologia da USP, São Paulo, Brazil John E. Randall ISSN 0945-9871 Bishop Museum, Honolulu, Hawaii, U.S.A. Publisher: Aquapress, Redazione aqua, I-27010 Miradolo Terme (Pavia), Italy Helen K. Larson www.aqua-aquapress.com Curator Emeritus, Fishes - Museum and Printer: Pronto Stampa Srl – Bergamo – Italy Art Gallery of the Northern Territory. Darwin, Copyediting and layout: Rossella Bulla Australia © 2013 aqua, International Journal of Ichthyology aqua, International Journal of Ichthyology

Description of a new species of annual fish of the genus Neofundulus (Cyprinodontiformes: Rivulidae) from the upper río Mamoré basin, Bolivia

Dalton Tavares Bressane Nielsen1 and Roger Brousseau2

1) Laboratório de Zoologia, departamento de Biologia, Universidade de Taubaté, Pça Marcelino Monteiro 63, CEP: 12030-010, Taubaté, SP, Brazil. E-mail: [email protected] 2) 8345 Bull Mountain Circle, Elk Grove, California 95758, USA. Email: [email protected]

Received: 03 March 2013 – Accepted: 17 June 2013 Abstract Résumé Neofundulus splendidus n. sp. is described from Bolivia, Neofundulus splendidus n. sp. est décrit, venant de Bolivie, departamento de Santa Cruz, río San Miguel, upper Departamento de Santa Cruz, rio San Miguel, bassin su - Mamoré basin. Neofundulus splendidus n. sp. is distin- périeur du Mamoré. Neofundulus splendidus n. sp. se distinguished from the remaining Neofundulus species by a com- gue des autres espèces de Neofundulus par une combinaison bination of characters: Large black spot at humeral region de caractéristiques: une grande tache noire dans la région with diameter of approximately 1.5-2.0 of pupil in males humérale avec un diamètre d’environ 1,5 - 2,0 de la pupille and females (vs. black spot in humeral region, when pre- pour les mâles et les femelles (contre une tache noire dans la sent, small- half of pupil), 40-41 scales in longitudinal région humérale, si elle est présente,, d’une petite moitié de series (vs. 35-38 in remaining congeners).This is the first la pupille), 40 - 41 écailles en séries longitudinales (contre record for the genus Neofundulus in río Mamoré basin. 35 - 38 pour les autres congénères). Il s’agit de la première oc currence du genre Neofundulus dans le bassin du rio Resumo Mamoré. Neofundulus splendidus n. sp. é desrcito da Bolívia, departamento de Santa Cruz, río San Miguel, alto bacia Sommario Mamoré. Neofundulus splendidus n. sp. destingue das Neofundulus splendidus n. sp. è descritto dalla Bolivia, demais espécies de Neofundulus pela seguintes combi- De par tamento de Santa Cruz, Rio San Miguel, bacino nações de características: Grande mancha preta na região superiore del Mamoré. Neofundulus splendidus n. sp. si dis- de humeral com diâmetro de aproximadamente 1.5-2.0 do tingue dalle altre specie di Neofundulus per una combi- diâmetro da pupila em machos e fêmeas (vs. pequena ou nazione di caratteri: presenza sia in maschi sia in femmine ausente mancha preta na região de humeral), 40-41 esca- di una grande macchia nera in regione omerale con mas em séries longitudinais (vs. 35-38 nas demais espé- diametro di circa 1.5-2.0 volte quello della pupilla (vs. cies). Este é o primeiro registro para o gênero Neofundulus macchia nera nella regione omerale, quando presente, pic- na bacia do rio Mamoré. cola e di diametro metà della pupilla), 40-41 scaglie in serie longitudinale (vs. 35-38 nei congeneri). Il genere Zusammenfassung Neofundulus non era stato ancora segnalato nel bacino del Neofundulus splendidus sp. n. wird vom Fluss San Miguel Río Mamoré. aus dem Einzugsgebiet des Mamoré-Oberlaufs in Bolivien, Departamento Santa Cruz, beschrieben. Die neue Art unterscheidet sich von anderen Arten der Gattung Neo- INTRODUCTION fundulus durch eine Kombination folgender Merkmale: Even though the first Neofundulus species N. großer schwarzer Fleck im Humeralbereich mit etwa dem paraguayensis, was described over a hundred years 1,5- bis 2,0-fachen Durchmesser der Pupille bei Män- ago (Eigenmann & Kennedy, 1903), few species of nchen und Weibchen (im Gegensatz zu einem kleinen the genus are so far known, when compared with schwarzen Fleck im Humeralbereich, der – wenn über- other Rivulidae genera, and few studies have been haupt vorhanden – etwa dem halben Pupillendurchmesser entspricht), 40-41 Schuppen in Längsreihen (35-38 bei conducted on this genus. den übrigen Angehörigen der Gattung). Es handelt sich So far, only five Neofundulus species have been um einen Erstnachweis der Gattung Neofundulus im Rio- described: N. paraguayensis (Eigenmann & Marmoré-Einzugsgebiet. Kennedy, 1903) from the río Paraguay basin, N.

109 aqua vol. 19 no. 3 - 19 July 2013 Description of a new species of annual fish of the genus Neofundulus (Cyprinodontiformes: Rivulidae) from the upper río Mamoré basin, Bolivia ornatipinnis Myers 1935 from the río Paraguay Costa (2006). For vertebral counts the caudal com- basin, N. parvipinnis Costa, 1988 from the rio pound centrum was counted as a single element. Cuiabá basin (río Paraguay basin), N. guaporensis Institutional abbreviations follow Sabaj-Pérez Costa 1988 from the rio Guaporé basin, and N. (2010), with addition of UNITAU (Universidade acutirostratus Costa 1992 from the rio das Velhas, de Taubaté), MNKP- Museo Noel Kempff, Santa rio São Francisco basin. Cruz de La Sierra, Bolivia. Osteological features Myers (1924) erected Neofundulus for Fundulus included in the description are those considered paraguayensis, which has as its type locality a pool phylogenetically informative in studies on Neofun- near the arroyo Trementina Paraguay. Aramburu et dulus (1988, 1992). al. (1962) studied material of the genus collected in Argentina and verified through meristic analysis and ray fin counts that Neofundulus paraguayensis Neofundulus splendidus, n. sp. could not be distinguished from Neofundulus (Figs 1-2, Table I) ornatipinnis, which they considered to be a syn- onym of the earlier. Holotype: MNKP-11205, 01 male, 63.5mm SL, Costa (1988) revalidated the two species, based Bolivia. Deposited at Museo Noel Kempff, road on number of rays of dorsal fin, number of scales San Miguel to San Juan, Departamento Santa in transversal series and caudal peduncle and in the Cruz, San Miguel city, río San Miguel, río Mamoré form of the fins he considered N. ornatipinnis e N. basin (16°46’22.0”S 61°11’12.6”W), altitude 440 paraguayensis separate species and described two m, 20 February 2012, Didier Pillet, Bruno Accorsi, additional species Neofundulus parvipinnis and Jean Marc Beltramon, Michel Beuchey& Christine Neofundulus guaporensis. Lambert. The last Neofundulus species of the genus to be Paratypes: MNKP-11206, five males 33.9-40.5 described, N. acutirostratus, was based on a single mm SL and six females, 33.5-37.1 mm SL, Bolivia. specimen collected in 1942 by A. L. Carvalho and Deposited at Museo Noel Kempff, Santa Cruz de G. S. Myers (Costa, 1992). La Sierra, ZUEC-7303, one male, 35.7 mm SL The genus Neofundulus is one of the several annual and one female, 36.2 mm SL. Deposited at Museu genera of Rivulidae, that is, they are considered de Zoologia da Universidade de Campinas, Camp- annual fish for living in temporary habitats and inas, São Paulo, Brazil, collected with the holotype. resisting desiccation by laying their eggs on the sub- Diagnosis: Neofundulus splendidus differs from stratum. Annual fish are unique in the fact that they the remaining species of the Neofundulus genus by develop their eggs very slowly and can go up through the male body with large black spot at humeral to three diapauses, depending on genetic and envi- region with diameter of approximately 1.5-2.0 of ronmental factors (Myers 1952; Wourms 1972). pupil in males and females (vs. black spot in The new species described below was found in a humeral region, when present, small), 40-41 scales roadside annual pool between the towns of San in longitudinal series (vs. 35-38 in remaining con- Miguel and San Juan, departamento de Santa geners), by color pattern, except N. guaporensis and Cruz, in the area near the río San Miguel, a tribu- N. acutirostratus, consisting in three to four yellow tary of the right bank of the río Mamore, Bolivia. lines in the body, alternated with two to three slender red lines, that become thicker and dark towards MATERIAL AND METHODS the caudal peduncle (vs. yellow lines lacking in N. Measurements were taken point-to-point under a paraguayensis, a single yellow line in N. ornatipin- stereomicroscope with a digital caliper to the near- nis, three yellow lines alternated with three red est 0.1 mm on the left side of the specimen follow- lines in N. parvipinnis), the color of pectoral fin in ing Costa (1995; 2007). Measurements are males, with finely delicate drawing with narrow expressed as percents of standard length (SL), white grading to yellow lines within a dark orange except the measurement of the head, which are background (vs. pectoral fin with black spots in recorded as percents of head length (HL). remaining congeners), presence of four to five Counts of vertebrae and pleural ribs were taken transverse black bars in dorsal fin in males (vs. from radiographs of the holotype, five males para- black bars absent or presence of small black spots, types and six females paratypes. Terminology for or else combinations of two black bars with small frontal squamation follows Hoedeman (1958) and black spots in the remaining Neofundulus species), aqua vol. 19 no. 3 - 19 July 2013 110 Dalton Tavares Bressane Nielsen and Roger Brousseau

Table I. Morphometric and meristic data for the holotype (H) and paratypes of Neofundulus splendidus.

H Paratypes Male Male n = 5 Females n = 6

Standard length (mm) 63.5 33.9-40.5 33.5-37.1 Percents of standard length Body depth 66.4 22.1-23.6 18.5-20.5 Caudal-peduncle depth 14.3 13.6-14.5 12.7-14.4 Pre-dorsal length 67.1 73.7-77.0 71.4-75.4 Pre-pelvic length 53.5 52.3-54.0 53.7-57.0 Length of dorsal-fin base 17.8 10.8-13.3 12.7-15.7 Length of anal-fin base 21.2 17.7-23.9 17.1-22.3 Caudal-fin length 18.4 24.2-28.4 22.4-30.4 Pectoral-fin length 21.4 19.8-24.1 15.7-26.9 Pelvic-fin length 11.8 11.3-17.9 11.6-16.1 Head length 24.7 29.3-32.4 28.4-33.9 Percents of head length Head depth 68.0 57.3-64.1 52.6-62.3 Head width 34.6 32.5-40.7 31.3-35.9 Lower jaw length 17.2 16.9-20.8 17.1-19.4 Eye diameter 28.0 27.9-29.2 26.5-29.4 Counts Dorsal fin 10 9-10 11-12 Caudal fin 22 22 22-23 Anal fin 14 14-15 15-16 Pelvic fin 07 07 6-7 Pectoral fin 14 12-14 12-14 Meristic Scales in longitudinal series 40 40-41 40-41 Scales in transversal series 09 09 09 Horizontal scales around 16 16 16 caudal peduncle

Fig. 1. Neofundulus splendidus, male, holotype, 63.5 mm SL: Bolivia, Departamento de Santa Cruz, San Miguel. Photo by G. Dethu.

111 aqua vol. 19 no. 3 - 19 July 2013 Description of a new species of annual fish of the genus Neofundulus (Cyprinodontiformes: Rivulidae) from the upper río Mamoré basin, Bolivia three transverse brown bars in the body of females pleural ribs of 17th and 19th vertebrae. Dorsal fin (vs. bars lacking in females of the remaining con- rays 10-13; anal fin rays 14-16; caudal fin rays 22- geners), transverse brown bars in dorsal and anal 24; pectoral fin rays 12-14; pelvic fin rays 7. fins in females (vs. brown bars absent or presence Scales large, cycloid. Body and head entirely of small black spots in dorsal fin of the females of scaled, except antero-ventral surface of head. Body remaining Neofundulus species). scales extending over caudal-fin base; no scales on Description: Morphometric and meristic data dorsal and anal fins. Frontal squamation E-pat- presented in Table I. Largest specimen examined terned; E-scales not overlapping medially; scales 63.5mm SL. Body elongated, slender, subcylindri- arranged in irregular circular pattern around A- cal, deeper than wide, compressed posteriorly. scale without exposed margins. Longitudinal series Greatest body depth at level of pelvic-fin base. of scales 40-41; transverse series of scales 9; scale Dorsal profile gently convex from snout to end of rows around caudal peduncle 16. Total vertebrae dorsal-fin base, approximately straight to slightly 34-35. Contact organs absent. concave on caudal peduncle. Ventral profile Coloration in life (Figs 1-2): slightly convex on head, almost straight from ante- Males: Side of body with yellow, red and irides- rior portion of abdominal region to end of anal-fin cent clear blue spots longitudinally aligned, base, slightly concave along caudal peduncle. becoming olive-green on the back, abdominal area Greatest body depth at level of pelvic-fin base. whitish. Large black spot on humeral region with Snout very blunt. Jaws long. diameter of approximately 1.5-2.0 of pupil. Sides Tip of dorsal and anal-fins slightly pointed in of head olive-green. Iris yellow, with dark, vertical male, rounded in female, lacking filamentous rays. brown bar crossing center of eye. Dorsal fin pale Anal fin approximately rectangular in males, longer yellow, with four to five transverse dark bars. Anal- than deep, reaching vertical through base of caudal fin dark olive-green, large golden yellow bar sur- fin. Anal-fin rounded, short, and without filamen- rounded by dark pigmentation near the basis, in tous rays in females. some specimens seemingly to continue into the Caudal-fin rounded to subtruncated in males and caudal-fin. Caudal-fin dark gray, with white and females, without filamentous rays. Pectoral-fin ellip- yellow spots irregularly distributed, and distal por- tical. Pelvic-fin pointed, tip reaching between base tion of the fin black. Pelvic fin dark orange, with- of 3rd and 4th anal-fin rays in male and 1st and 2nd out spots or lines. Pectoral fin with finely delicate anal-fin rays in female. Pelvic-fin bases in close prox- drawing composed by narrow white grading to yel- imity in males, and separated by small interspace, in low lines within a dark orange background. females. Dorsal-fin origin on vertical through base Females: Side of body light brown, with three of 5th or 6th anal-fin ray, and between neural spines longitudinal brown stripes. Large black spot on of 22nd and 24th vertebrae. Anal-fin origin between humeral region with diameter of approximately

Fig. 2. Neofundulus splendidus, female, paratype, 37.1 mm SL: Bolivia, Departamento de Santa Cruz, San Miguel. Photo by G. Dethu. aqua vol. 19 no. 3 - 19 July 2013 112 Dalton Tavares Bressane Nielsen and Roger Brousseau

1.5-2.0 of pupil. Dorsum light brown, abdominal tary of the rio Madeira basin, the rio Guaporé region whitish. Sides of head grey, with a pale basin, the new species extends the distribution of metallic yellow tinge on opercle. Iris yellow, with the genus and suggests that the genus might be pale brown horizontal bar crossing center of eye. more widespread in other hydrographic basins on Dorsal and anal fins hyaline, with longitudinal the right bank of the Amazon basin. brown bars. Caudal-fin pale yellow, with transverse Other species of annual fish of several genera, narrow, light brown bars. Pectoral fin yellow with Papiliolebias, Spectrolebias, Trigonectes, Moema, small brown spots. Pelvic fin pale yellow. Aphyolebias, and Pterolebias were found recently in Distribution: Neofundulus splendidus is currently Bolivia, at the río Mamore, rio Guaporé and río only known from its type-locality. Paraguay basins (Costa 1996, 1998a, Costa et al. Habitat (Fig. 3): The type-locality lies at the 1996, Costa et al. 1997). Since the area searched plateau area (440 ma.s.l.), just southeast of Llanos was relatively small when the total area of these de Mojos, which occupy much of the río Mamoré hydrographic basins is considered, and also consid- basin in Bolivia (see Loubens et al., 1992). The ering the remoteness of most of the area, it is very pool is located beside the highway, between the likely that further fieldwork will discover addi- cities of San Miguel to San Juan. Water tempera- tional new rivulid species in this region. ture of the pool was 24.6°C, at the depth of 40 cm. Neofundulus is distinguished from the remaining In the marginal area of the pool, at the depth of 10 genera of Rivulidae family by having the following cm, water temperature was 29°C. Neofundulus synapomorphies: fourth pectoral radial not splendidus was the only fish species present in the expanded ventrally, a subasal longitudinal white- pool, where it was found over its entire area. The yellowish stripe on the anal fin in males, transverse pool presented clean water, pH 6.4, 0.5 mg/l of black bars on caudal fin of females, and black spots dissolved iron (Fe) and electric conductivity 67 µS. on pectoral fin of males (Costa 1998b) Neofundu- Other animals recorded at the pool were Phylome- lus splendidus presents very narrow yellow lines on dusa sp. tadpoles, clams and freshwaters crabs. A the pectoral fins, instead of black spots present on dense coverage of aquatic vegetation mainly the other Neofundulus species, thus an autapomor- formed by Echinodorus sp. and Nymphaea sp., was phy of this species. Neofundulus splendidus appears present at the pool. to be closer to Neofundulus parvipinnis, since Behavior in captivity: The male initiates the because both species share 14 to 16 anal-fin rays courtship expanding its unpaired fins next to the and a color pattern of the males presenting longi- female, forming with their bodies, angles of 10° or tudinal yellow stripes. 15°. Sometimes the male touch, with its mouth, the female abdominal region. After the male dis- play, it touches the substratum with its snout, and the couple dives in the substratum. The female lay two or three large-sized (1.5 mm diameter) eggs per day. Males can be sometimes aggressive towards the female, which makes necessary to separate the couple after the end of courtship. Etymology: In reference to the exuberant colors of the male. An adjective.

DISCUSSION Neofundulus splendidus can be easily identified from its congeners by the color pattern of the pectoral-fin in males, and by the presence of a large black spot in the humeral region of males and females. Neofundulus was previously found in southern Bolivia in the Paraguay basin (Costa et al. 1996), but this is the first record of the genus for Fig. 3. Type locality of Neofundulus splendidus near río the río Mamore basin. Although the genus Neo- San Miguel, upper río Mamoré basin, Bolivia. Photo by fundulus had already been found in another tribu- D. Pillet.

113 aqua vol. 19 no. 3 - 19 July 2013 Description of a new species of annual fish of the genus Neofundulus (Cyprinodontiformes: Rivulidae) from the upper río Mamoré basin, Bolivia

Key to Neofundulus species based on Costa 1992, COSTA, W. J. E. M. 1996. A new Specie of the annual fish with the addition of Neofundulus splendidus: genus Pterolebias (Cyprinodontiformes: Rivulidae) from the Rio Mamoré basin, Bolivian Amazon. Neotropical 1a. Origin of dorsal fin on vertical through base of Ichthyology, 7: 91-96. COSTA, W. J. E. M. 1998a. Revision of the Neotropical first or second anal-fin rays ...... 2 annual fish genus Plesiolebias (Cyprinodontiformes: 1b. Origin of dorsal fin on vertical through base of Rivulidae). Ichthyological Exploration of Freshwaters, 8: fifth or sixth anal-fin rays...... 4 313-334. 2a. Caudal-fin rounded; tip of pelvic fin in males COSTA, W. J. E. M. 1998b. Phylogeny and classification of reaching fourth to sixth anal-fin ray;16 hori- Rivulidae revisited: Origin and evolution of annualism zontal scales around caudal peduncle ...... and miniaturization in rivulid fishes (Cyprinodonti- ...... N. paraguayensis formes: Aplocheiloidei). Journal of Comparative Biology, 2b. Caudal-fin truncated; tip of pelvic fin in males 3: 33-92. COSTA, W. J. E. M. 2006. Descriptive morphology and reaching first or second ray of anal fin; 18 to 20 phylogenetic relationship among species of the Neotrop- horizontal scales around caudal peduncle ...... ical annual killifish genera Nematolebias and Simp- ...... 3 sonichthys (Cyprinodontiformes: Aplocheiloidei: Rivuli- 3a. Lateral profile of the head pointed; caudal dae). Neotropical Ichthyology, 4: 1-26. peduncle depth 17.0 % SL...... COSTA, W. J. E. M. 2007. Taxonomic revision of the sea- ...... N. acutirostratus sonal South American killifish genus Simpsonichthys 3b. Lateral profile of the head rounded; caudal (Teleostei: Cyprinodontiformes: Aplocheiloidei: Rivuli- peduncle depth 12.5-15.0 % SL ...... dae). Zootaxa, 1669: 1-134. COSTA, W. J. E. M., BARRERA, S. & SARMIENTO, J. 1996...... N. ornatipinnis A new species of annual fish genus Pterolebias (Cyprino - 4a. 12 rays anal fin...... N. guaporensis dontiformes: Rivulidae) from the Rio Mamoré basin, 4b. 14 to 16 rays anal fin ...... 5 Bolivian Amazon. Ichthyological Exploration of Freshwa- 5a. 35 to 36 scales in longitudinal series ...... ters, 7: 91-95...... N. parvipinnis COSTA, W. J. E. M., BARRERA, S. & SARMIENTO, J. 1997. 5b. 40 to 41 scales in longitudinal series ...... Simpsonichthys filamentosus, une nouvelle espèce des Lla- ...... N. splendidus nuras Benianas, bassin du Rio Mamoré, Bolivia. Revue Française d’Aquariologie, 24: 83-86. EIGENMANN, C. H. & KENNEDY, C. H. 1903. On a col- ACKNOWLEDGEMENTS lection of fishes from Paraguay with a synopsis of the We are grateful to Didier Pillet, Bruno Accorsi, American genera of cichlids. Proc. Acad. Nat. Sci. Philad., Jean Marc Beltramon, Michel Beuchey, and Chris- 477-537. tine Lambert for senting the type-material, Itamar HOEDEMAN, J. J. 1958.The frontal scalation pattern in Martins for support at the laboratory, Flávio C. T. some groups of tooth carps. Bulletin of Aquatic Biology, 1: Lima for useful suggestions to the manuscript, 23-28. Guillaume Dethu for the pictures and Didier Pillet MYERS, G. S. 1924. A new poeciliid fish from Congo, with for informations about breeding behavior. remarks on funduline genera. Amer. Mus. Novitates, 116: 1-9. MYERS, G. S. 1935. Four new freshwater fishes from Brzil, REFERENCES Venezuela, and Paraguay. Proc. Biol. Soc. Wash., 48: 7-14. ARÁMBURU, A. S. A., ARÁMBURU, R. H. & RINGUELET, R. MYERS, G. S. 1952. Annual fishes. Aquarium Journal, 2: A. 1962. Peces paranaenses nuevos para la fauna 125-141. argentina. Physis XXIII, 65: 223-239. NIELSEN, D. T. B. 2008. Simpsonichthys e Nematolebias. COSTA, W. J. E. M. 1988. Sistemática e distribuição do Cabral Editora e Livraria Universitária, Taubaté. 235 pp. gênero Neofundulus (Cyprinodontiformes: Rivulidae), SABAJ-PÉREZ, M. H. 2010. Standard symbolic codes for com a descrição de duas espécies novas. Rev. Brasil. Biol., institutional resource collections in herpetology and 48 (2): 103-111. ichthyology: an Online Reference. Verson 1.5 (4 Oct COSTA, W. J. E. M. 1992. Descrição de uma nova espécie 2010). Electronically accessible at http://www.asih.org/, de Neofundulus (Cyprinodontiformes: Rivulidae), da American Society of Ichthyologists and Herpetologists, bacia do rio São Francisco, Brasil. Revista Brasileira de Washington, DC. Biologia 52 (4): 615-618. WOURMS, J. P. 1972. The developmental biology of annual COSTA, W. J. E. M. 1995. Pearl killifishes: The Cynolebi- fishes III. Pre-embryonic and embryonic diapause of atinae. Systematics and Biogeography of a Neotropical variable duration in the eggs of annual fishes. Journal of annual fish subfamily (Cyprinodontiformes: Rivulidae). Experimental Zoology, 182: 389-414. TFH, Neptune City, 128 pp.

aqua vol. 19 no. 3 - 19 July 2013 114 aqua, International Journal of Ichthyology

Spectrolebias pilleti, a new annual Killifish (Cyprinodontiformes: Rivulidae: Cynolebiatinae) from the upper río Mamoré basin, Bolivia

Dalton Tavares Bressane Nielsen1 and Roger Brousseau2

Received: 29 March 2013 – Accepted: 17 June 2013

Abstract posição da nadadeira dorsal, pois possui sua origem ante- Spectrolebias pilleti n. sp. is described from a temporary rior a origem da nadadeira anal. pool from the upper río Mamoré basin, Departamento Santa Cruz, Quimone city, Bolivia. The new species is dis- Zusammenfassung tinguished from all congeners by the unique color pattern Spectrolebias pilleti n. sp. wird von einem zeitweiligen of males:3 to 5 vertical blue bars alternating with a light Tümpel im Einzugsgebiet des oberen Mamoré, Departa- reddish brown background in males (vs. absence of bars), mento Santa Cruz, in der Stadt Quimone, Bolivien, by the presence of an iridescent blue spot on the flank in beschrieben. Die Vertreter dieser neuen Art unterscheiden female (vs. presence of one or more black spots), by the sich von allen anderen Angehörigen der Gattung durch das absence of filaments at the tips of dorsal fin and anal fin in unverkennbare Farbmuster: 3 bis 5 senkrechte blaue males, except Spectrolebias reticulatus (vs. long filaments on Streifen, die sich von einem hell rötlich braunen Unter- dorsal and anal fins in S. brousseaui and S. filamentosus, fil- grund abheben bei den Männchen (bei den anderen Arten aments on the dorsal fins of S. semiocellatus and S. inae- fehlen Streifen), schillernd blauer Fleck an der Seite bei quipinnatus, or filaments in the anal fins of S. chacoensis) den Weibchen (im Gegensatz zu einem oder mehreren and by the presence of scales covering the base of anal fin schwarzen Flecken bei den anderen Arten), das Fehlen von in males (vs. absence of these scales in the remaining con- fädigen Strukturen an den Spitzen von Rücken- und After- geners). Also distinguished from all congeners, with the flosse der Männchen, außer bei Spectrolebias reticulatus exception of S. reticulatus, by the position of the dorsal fin, (S. brousseaui und S. filamentosus haben lange Filamente an which has its origin anterior to the origin of the anal fin in Rücken- und Afterflosse, S. semiocellatus und S. inaequi- males. pinnatus zeigen Filamente an der Rückenflosse, S. chacoensis an der Afterflosse) sowie Schuppen an der Basis der Resumo Afterflosse bei Männchen (die bei den anderen Arten der Spectrolebias pilleti n. sp é descrita de poças temporárias Gattung fehlen). Außerdem ist die Lage der Rückenflosse da bacia do alto rio Mamoré. Departamento de Santa im Vergleich zu allen anderen Angehörigen der Gattung Cruz, cidade de Quimone, Bolívia. A nova espécie se difer- außer S. reticulatus ein Unterscheidungsmerkmal; sie setzt encia das demais espécies do gênero por possui padrão de bei den männlichen Vertretern der neuen Art vor der coloração único nos machos: 3 à 5 barras verticais azuis Afterflosse an. alternadas com claras barras castanho no corpo dos machos (vs. ausência de barras)pela presença de uma mancha azul Résumé iridescente no flanco das fêmeas (vs. presença de uma ou Spectrolebias pilleti n. sp. est décrit en provenance d’une mais manchas pretas), pela ausência de filamentos nas pon- mare temporaire du bassin supérieur du rio Mamoré, tas das nadadeiras dorsal e anal (vs. longos filamentos nas Departamento Santa Cruz Quimone city, Bolivie. La nou- nadadeiras dorsal e anal em S. brousseaui e S. filamentosus, velle espèce se distingue de tous ses congénères par la filamentos na ponta da nadadeira dorsal em S. semiocella- couleur unique des mâles: 3 à 5 barres verticales bleues tus e S. inaequipinnatus e filamentos na ponta da nadadeira alternant sur un fond légèrement rouge brun pour les anal em S. chacoensis) e presença de escamas cobrindo a mâles (contre l’absence de barres), par une tache bleue iri- base da nadadeira anal em machos (vs. ausência de escamas descente sur le flanc de la femelle (contre une ou plusieurs nas demais espécies do gênero). Também se distingue das taches noires), par l’absence de filaments au bout de la dor- demais espécies do gênero, exceto S. reticulatus, pela sale et de l’anale pour le mâle, sauf pour Spectrolebias reti-

115 aqua vol. 19 no. 3 - 19 July 2013 Spectrolebias pilleti, a new annual Killifish (Cyprinodontiformes: Rivulidae: Cynolebiatinae) from the upper río Mamoré basin, Bolivia culatus (contre de longs filaments sur la dorsale et l’anale According to Costa (2006) Spectrolebias is distin- pour S. brousseaui et S. filamentosus, des filaments sur les guished from all other cynolebiasins by two unam- dorsales de S. semiocellatus et S. inaequipinnatus, ou des fil- biguous synapomorphies: a long (vs. short) aments sur l’anale de S. chacoensis ) et par des écailles cou- hyomandibula and a narrowed (vs. wide) proximal vrant la base de l’anale pour les mâles (contre l’absence des- dites écailles chez les autres congénères). L’espèce se dis- tip of the fourth ceratobranchial. tingue aussi de tous les congénères, à l’exception de S. reti - All species of the genus Spectrolebias are annual cu latus, par la position de la dorsale, dont la naissance est fish and live in temporary water pools where they antérieure à celle de l’anale pour les mâles. lay eggs by diving into the substrate. As the remaining representatives of the tribe Cynolebi- Sommario asini, species of the genus Spectrolebias possess an Spectrolebias pilleti n. sp. è descritto da una pozza tempo- annual life cycle. At the end of the rainy season the ranea dal bacino superiore del Rio Mamoré, Departa- pool dries and all fish die, but their eggs survive mento Santa Cruz, Quimone, Bolivia. La nuova specie si and develop throughout the dry season in the sub- di stingue da tutte le congeneri per il colore unico della livrea maschile, caratterizzata da 3-5 barre blu verticali su stratum. As soon as the rain starts, the pools fill up un fondo marrone rossastro (vs. assenza di barre), dalla with water and the eggs hatch, starting a new gen- presenza negli individui femmina di una macchia blu iri- eration that will on his turn grow, breed and keep descente sul fianco (vs. presenza di uno o più punti neri), the cycle going (Myers, 1952; Costa, 1995; per l'assenza di filamenti sulle punte sia della pinna dorsale Nielsen, 2008). sia della pinna anale nei maschi, tranne Spectrolebias reticu- The present contribution describes an additional latus (vs. lunghi filamenti sulle pinne dorsale e anale in species of Spectrolebias from Bolivia, the third S. brousseaui e S. filamentosus, filamenti sulle pinne dorsali species known for the country. This new species in S. semiocellatus e S. inaequipinnatus, o filamenti nelle pinne anali in S. chacoensis) e dalla presenza di scaglie che was found in the basin of the río San Pablo, a trib- co pro no la base della pinna anale nei maschi (vs. assenza di utary on the right side of the río Mamoré, Qui- queste scale nei restanti congeneri). Si distingue anche da mone city, Departamento de Santa Cruz. It was tutte le specie congeneri, con l'eccezione di S. reticulatus, original discovered in 2011, at the same location of per la posizione della pinna dorsale, la cui origine nei the specimens collected in 2012, which served as maschi è anteriore all'origine della pinna anale. the basis for this work.

MATERIAL AND METHODS INTRODUCTION Measurements were taken point-to-point under a The species of the genus Spectrolebias Costa and stereomicroscope with a digital caliper to the near- Nielsen, 1997 is distributed across the tributaries est 0.1mm, on the left side of the specimens, when- of the right margin of Amazon River, with a single ever possible, following Costa (1995, 2007). Mea- exception, Spectrolebias chacoensis which is found in surements are expressed as percentages of standard the río Paraguay basin. Currently, the following length (SL), except subunits of the head, which are species are: S. chacoensis (Amato, 1986), from the recorded as percentages of head length (HL). río Paraguay basin, Paraguay and Argentina; S. In the description, counts of vertebrae and pleural costai (Lazara, 1991) from the rio Araguaia and rio ribs were taken from radiographs of the holotype Tocantins basins, Brazil; S. semiocellatus Costa & and two female paratypes. Terminology for frontal Nielsen, 1997, from the rio Araguaia basin, Brazil; squamation follows Hoedeman (1958) and Costa S. filamentosus (Costa, Barreira & Sarmiento, (2006). For vertebral counts the caudal compound 1997), from the río Madeira basin, Bolivia; S. retic- centrum was counted as a single element. Osteo- ulatus (Costa & Nielsen, 2003), from the rio logical features included in the description are Xingu basin, Brazil; S. inaequipinnatus (Costa & those considered phylogenetically informative by Brasil, 2008), from the rio Tocantins basin, Brazil, recent studies on Spectrolebias (Costa 2006, 2007, and S. brousseaui Nielsen, 2013, from the río 2010). Institutional abbreviations follow Sabaj- Mamoré basin, Bolivia. Pérez (2010), with addition of UNITAU (Univer- Spectrolebias was originally considered a genus sidade de Taubaté) and MNKP- Museo Noel (Costa & Nielsen, 1997). After a phylogenetic Kempff. Comparative material of other rivulids analysis made by Costa (2006), was considered one examined in the present study is listed in Costa of the five subgenera of Simpsonichthys, but latter, (2007). Costa (2010) promoted it back to a genus status. aqua vol. 19 no. 3 - 19 July 2013 116 Dalton Tavares Bressane Nielsen and Roger Brousseau Spectrolebias pilleti, n. sp. covering the basis of anal fin in males (vs. absence (Figs 1-2, Table I) of scales covering the basis of the anal fin in the remaining congeners). Distinguished from all con- Holotype: Deposited at Museo Noel Kempff, geners, except S. reticulatus, by the position of the Santa Cruz de La Sierra, Bolivia. (MNKP-11153) dorsal fin, which has its origin anterior to the ori- male 28.9 mm SL: Bolivia, Departamento Santa gin of the anal fin in males, and by possessing the Cruz, Quimone city, 45.40 km west of San José tips of both dorsal and anal-fins pointed in males, dos Chiquitos, temporary pool near the río Qui- without elongated rays (vs. tips of both dorsal and mone, the río San Pablo basin, (17°42’48.1”S anal-fins pointed in males, with elongated rays, in 61°08’56.1”W), altitude 263 m, 26 February the remaining congeners). 2012, Didier Pillet, Jean Marc Beltramon, Michel Description: Morphometric data presented in Beuchey & Christine Lambert. Table I. Largest specimen examined 30.0 mm SL. Paratypes: Deposited at Museo Noel Kempff, Dorsal profile slightly concave on head, convex Santa Cruz de La Sierra, Bolivia (MNKP-11154) from nape to end of dorsal-fin base, approximately five male and eight females (24.3-30.0 mm SL), straight along caudal peduncle. Ventral profile collected with the holotype. Deposited at Universi- slightly convex from tip of jaw to origin of caudal dade de Campinas, Campinas, São Paulo, Brazil. peduncle and slightly concave from latter point to (ZUEC 7296), one male (36.0 mm SL), two uppermost procurrent caudal-fin rays. Body mod- females (22.5-26.0 mm SL), collected with the erately deep, compressed, greatest body depth on holotype. vertical slightly ahead to anal-fin origin. Eye posi- Diagnosis: The new species differs from the tioned on lateral portion of side of head. Snout remaining representatives of the genus Spectrolebias blunt. Large number of labial papillae in males. by the unique color pattern of males: 3 to 5 verti- Urogenital papilla cylindrical and long in males, cal blue bars alternating within a light reddish pocket-shaped in females. Tip of both dorsal and brown background in males (vs. absence of bars), anal-fins pointed in males without elongated rays; by the presence of a iridescent blue spot on the dorsal fin in females rounded to slightly pointed flanks of the females (vs. presence of one or more without filaments; anal-fin rounded in females. black spots in females) and by the presence of scales Dorsal-fin rays unbranched. Caudal-fin rounded.

Fig. 1. Spectrolebias pilleti, male, holotype, 28.9mm SL. MNKP-11153. Photo by G. Dethu.

117 aqua vol. 19 no. 3 - 19 July 2013 Spectrolebias pilleti, a new annual Killifish (Cyprinodontiformes: Rivulidae: Cynolebiatinae) from the upper río Mamoré basin, Bolivia

Table I. Morphometric and meristic data for the holotype (H) and paratypes of Spectrolebias pilleti.

H Paratypes Male Male n = 6 Females n = 8 Standard length (mm) 28.9 27.6-30.0 22.2-24.3 Percents of standard length Body depth 34.1 33.8-36.1 31.7-35.6 Caudal peduncle depth 11.2 12.7-14.0 11.5-13.5 Pre-dorsal length 45.8 43.9-44.6 50.2-59.7 Pre-pelvic length 42.4 39.5-42.4 43.7-50.1 Length of dorsal-fin base 38.9 34.1-39.3 21.1-24.5 Length of anal-fin base 39.2 35.1-40.8 23.1-27.4 Caudal-fin length 26.8 25.0-28.4 22.2-27.1 Pectoral-fin length 29.7 28.2-34.2 18.9-26.4 Pelvic-fin length 10.9 9.5-13.7 8.5-11.2 Head length 29.7 28.7-30.6 28.6-31.5 Percents of head length Head depth 86.3 86.1-95.3 92.2-98.6 Head width 40.9 38.5-43.2 42.6-48.5 Snout length 6.9 6.8-7.2 14.3-14.8 Lower jaw length 11.6 11.6-13.2 14.1-15.7 Eye diameter 34.4 28.7-34.4 27.5-32.8 Counts Dorsal fin 23 23-24 20-21 Caudal fin 22 20-22 19-21 Anal fin 24 22-24 21-22 Pelvic fin 5 5 5 Pectoral fin 11 10-11 8-9 Scales in longitudinal series 28 29 Scales in transversal series 12 9 Horizontal scales around caudal peduncle 10 16

Fig. 2. Spectrolebias pilleti, female, paratype, 24.3 mm SL. MNKP-11154. Photo by G. Dethu. aqua vol. 19 no. 3 - 19 July 2013 118 Dalton Tavares Bressane Nielsen and Roger Brousseau

Dorsal-fin origin on vertical anterior to anal-fin Coloration in life (Figs. 1-2): origin in males, anal-fin origin on vertical between Males: Head and trunk to the vertical line passing base of 1st and 3th dorsal-fin rays; dorsal-fin origin through pelvic-fin origin light brown. Posterior two- on vertical through base of 2nd anal-fin rays in thirds of body with three to five blue bars, over the females. Tip of each pelvic-fin reaching base of 8rd light-brown background. including the unpaired to 9th anal-fin rays in males, and base of 1st to 2nd fins. Light blue spots sparsely distributed across the anal-fin rays in females. Dorsal-fin origin between caudal peduncle. Color pattern can be assymetrical; neural spines of fifth and sixth vertebrae in males, a single male can have three vertical blue bars in one and between neural spines of tenth and eleventh side of the body and five bars in the other side. vertebrae in females. Anal-fin origin at twelfth Tip of jaw darkened. Dorsal and anal-fins blue, pleural ribs of vertebrae in males, and pleural ribs with irregularly-distributed small light blue spots. of vertebrae ninth in females. Caudal-fin blue with higher concentration of light Dorsal-fin rays 19-20 in males, 20-21 in females, blue spots than dorsal and anal fins. Hyaline pec- anal-fin rays 22-24 in males, 21-22 in females, cau- toral fin. Pelvic-fin blue without light blue spots. dal-fin rays 20-21, pectoral-fin rays 10-11 and pelvic Black bar vertically crossing the eye. fin-rays 5. Frontal squamation E-patterned; Females: Side of body light brown, with 6-8 light- E-scales overlapping medially; no scale anterior to gray bars; one blue iridescent single spots on mid- G-scale; supraorbital scales absent. Longitudinal body. Scattered iridescent blue scales present in gray series of scales 27-28; transverse series of scales 12-13; bars. Sides of head light brown, opercular region scale rows around caudal peduncle 9-10. Contact pale greenish golden. Iris light yellow, with dark organ absent on flanks and pectoral fins in males. vertical brown bar across center of eye. Fins hyaline.

Fig. 3. Type locality of Spectrolebias pilleti near of Quimone city, Departamento de Santa Cruz, Bolivia. Photo by D. Nielsen.

119 aqua vol. 19 no. 3 - 19 July 2013 Spectrolebias pilleti, a new annual Killifish (Cyprinodontiformes: Rivulidae: Cynolebiatinae) from the upper río Mamoré basin, Bolivia

Distribution (Fig. 3): Only known from the type- tions about 60 cm. In the marginal area of the locality, a pool near of the río Quimone, tributary pool, at the depth of 10 cm, water temperature was of the río San Pablo, tributary of the río Mamoré, 31°C. Specimens of Spectrolebias pilleti were col- Departamento Santa Cruz, Quimone city, Bolivia. lected in depths no shallower than 20 cm depth. Habitat (Fig. 4): The type-locality lies at the The pool presented clean water, pH 6.8, dissolved plateau area (263 m.a.s.l.), at the border of the iron (Fe) 0,25 mg/l and electric conductibility 100 Llanos de Chiquitos, to the north of the Bañados µS. Other fish species collected syntopically were del Izozog, where several rivers of the upper río exclusively annual fishes: Spectrolebias brousseaui, Mamoré basin drain, eastern Bolivia (see Loubens Neofundulus sp., and Papiliolebias sp. Other ani- et al., 1992). The annual pool is very large, with mals recorded were Phylomedusa sp. tadpoles, most of its area within a dense, difficult to access clams and freshwaters crabs. The aquatic vegeta- forest. The temperature on the water surface was tion was very dense, composed by Echinodorus sp., 28°C and, in the deepest portion and banks. Aver- Utricularia sp. and Nymphaea sp. age depth of the pool is 40 cm, with deepest por- Etymology: The specific name is in honor of

Fig. 4. Geographic distribution of members Spectrolebias. Photo by D. Nielsen aqua vol. 19 no. 3 - 19 July 2013 120 Dalton Tavares Bressane Nielsen and Roger Brousseau environmentalist Didier Pillet, one of the collectors The limited amount of research along with the of the new species. difficulty of accessing this area (primarily due to the small number of roads in the region), raises the DISCUSSION possibility that many other annual fish species Albeit being still little explored, field exploration could be found in the Bolivian part of the Amazon during the last few years in Bolivia has revealed a basin. Members of the Spectrolebias genus are cur- large number of annual species, besides Spec- rently found in the Araguaia, Tocantins e Xingu trolebias, annual fishes from the genera Papili- basins in Brazil and in the río Mamoré basin in olebias, Trigonectes, Neofundulus, Moema, Aphy- Bolivia. Additionally, between those river basins olebias, and Pterolebias (Costa 1996, 1998, Costa, (all major southern tributaries of the Amazon Barrera, & Sarmiento 1997), were recorded for the River) are the rio Guaporé and rio Tapajós, and it country, though most of these records are still is plausible that Spectrolebias species might occur in unpublished (see Nielsen, 2013). There are also these intervening river basins. non annual Rivulidae species, from the genera The color pattern of the body, with the alternated Cynodontichthys and Anablepsoides. Such high blue and reddish brown bars, the absence of fila- diversity is probably a result of a combination of ments at the tip of anal and dorsal fins, and the factors, i.e., favorable geography, vegetation, and small iridescent blue single spots present in females high river drainage density and high precipitations body are autapomorphies of Spectrolebias pilleti level across this region, all favorable to the mainte- within the genus Spectrolebias. All three species of nance dispersion of annual fishes. The possibility Spectrolebias occurring in Bolivia (S. pilleti, S. fila- of finding new species in this region was raised in mentosus, and S. brousseaui) can be found in the río Nielsen 2013. The discovery of S. pilleti reinforces San Pablo basin, a tributary of the río Mamoré, this idea. suggesting that additional species of Spectrolebias Males of Spectrolebias pilleti differ from S. can be expected to occur in the río Mamoré basin brousseaui by the absence of dark blue coloration given its large area, little coverage by field research and bright blue spots aligned along the body of and difficult access. males (vs. presence), by the smaller size of bright blue spots on unpaired fins (vs. larger spots, half ACKNOWLEDGEMENTS the diameter of the pupil), by having pelvic-fin Thanks are due to Itamar Alves Martins from bases in contact (vs. pelvic-fin bases separated by Universidade de Taubaté (UNITAU) for laboratory interspace), pelvic-fin dark blue lacking spots (vs. support and to Guillaume Dethu for help with pic- presence of iridescent blue spots in the center), tures. André Carletto and Flávio C. T. Lima absence of contact organs on flank scales in males (ZUEC) reviewed the manuscript and offered use- (vs. presence), tip of pelvic-fin reaching the 8th or ful suggestions. 9th anal-fin ray (vs. 3rd to 5th), lower predorsal length (43.9-44.6% SL vs. 47.2-51.5%), lower caudal-fin length (25.0-28.4% SL vs. 29.6-32.6%), REFERENCES longer pectoral-fin length (28.2-34.2% SL vs. 21.1- AMATO, L. H. 1986. Seis espécies nuevas Del gênero 22.9%), lower head width (38.5-43.2% HL vs. Cynolebias Steindachner, 1876, de Uruguay y Paraguay (Cyprinodontiformes, Rivulidae). Comunicaciones Zoo- 54.6-55.6%), greater number of dorsal-fin rays (23- logicasdel Museo de Historia Natural de Montevideo 11 24 vs. 21-22), fewer number of pectoral fin (10-11 (162): 1-27. vs. 12), greater number of scales in transverse COSTA, W. J. E. M. 1995. Pearl killifishes: The Cynolebi- series(12-13 vs. 8-9), and fewer number of horizon- atinae. Systematics and Biogeography of a Neotropical tal scales around caudal peduncle ( 9-10 vs. 14-16). annual fish subfamily (Cyprinodontiformes: Rivulidae). Females of S. pilleti differ from females TFH Publications, Inc., Neptune City, 128 p. of S. brousseaui by a longer head depth (92.2-98.6 COSTA, W. J. E. M. 1996. A new Specie of the annual fish % HL vs. 78.1-91.6), lower head width (42.6- genus Pterolebias (Cyprinodontiformes: Rivulidae) from the Rio Mamoré basin, Bolivian Amazon. Neotropical 48.5% HL vs. 53.0-59.6), lower snout length Ichthyology, 7: 91-96. (14.3-14.8% HL vs. 18.7-23.4), greater number of COSTA, W. J. E. M. 1998. Revision of the Neotropical dorsal-fin rays (20 vs. 15-16), fewer number of annual fish genus Plesiolebias (Cyprinodontiformes: pectoral fin-rays ( 8-9 vs. 10-11), and greater num- Rivulidae). Ichthyological Exploration of Freshwaters, 8: ber of scales in transverse series (12-13 vs. 10). 313-334.

121 aqua vol. 19 no. 3 - 19 July 2013 Spectrolebias pilleti, a new annual Killifish (Cyprinodontiformes: Rivulidae: Cynolebiatinae) from the upper río Mamoré basin, Bolivia

COSTA, W. J. E. M. 2006. Descriptive morphology and HOEDEMAN, J. J. 1958.The frontal scalation pattern in phylogenetic relationship among species of the Neotrop- some groups of tooth carps. Bulletin of Aquatic Biology, ical annual killifish genera Nematolebias and Simp- 1: 23-28. sonichthys (Cyprinodontiformes: Aplocheiloidei: Rivuli- LAZARA, 1991. Cynolebias lacortei, Cynolebias costai, and dae). Neotropical Ichthyology, 4: 1-26. Cynolebias aruana: three new species of cloud fish from COSTA, W. J. E. M. 2007. Taxonomic revision of the sea- Brazil (Teleostei, Cyprinodontiformes, Rivulidae). Jour- sonal South American killifish genus Simpsonichthys nal of the American Killifish Association, 23: 139-152. (Teleostei: Cyprinodontiformes: Aplocheiloidei: Rivuli- LOUBENS, G. L., LAUZANNE & LE GUENNEC, B. 1992. dae). Zootaxa, 1669: 1-134. Lesmilieuxaquatiques de larégion de Trinidad (Beni, COSTA, W. J. E. M. 2010. Historical biogeography of Amazonie bolivienne). Revue d’Hydrobiologie Tropical cynolebiasine annual killifishes inferred from dispersal- 25 (1): 3-21. vicariance analysi. Journal of Biogeography. (on-line; doi: MYERS, G. S. 1952. Annual fishes. Aquarium Journal, 10.1111/j.1365-2699.2010.02339.x). 23: 125-141. COSTA, W. J. E. M. & NIELSEN, D. T. B. 1997. A new NIELSEN, D. T. B. 2008. Simpsonichthys e Nematolebias. genus and species of annual fish (Cyprinodontiformes: Cabral Editora e Livraria Universitária, Taubaté. 235pp. Rivulidae) from the Araguaia basin, central Brazil. Ichthy- Nielsen, D. T. B. 2013. Spectrolebias brousseaui (Cyprin- ological Exploration of Freshwaters, 7: 257-265. odontiformes: Rivulidae: Cynolebiatinae), a new annual COSTA, W. J. E. M., BARRERA, S. & SARMIENTO, J. 1997. fish from the upper río Mamoré basin, Bolivia. Neotropi- Simpsonichthys filamentosus, une nouvelle espècedes Lla- cal Ichthyological. 11 (1): 81-84. nuras Benianas, bassin du río Mamoré, Bolivia. Revue SABAJ-PÉREZ, M. H. 2010. Standard symbolic codes for Française d’Aquariologie, 24: 83-86. institutional resource collections in herpetology and COSTA, W. J. E. M. & NIELSEN, D. T. B. 2003. Simp- ichthyology: an Online Reference. Verson 1.5 (4 Oct sonichthys reticulatus n. sp. (Cyprinodontiformes: Rivuli- 2010). Electronically accessible at http://www.asih.org/, dae) a new annual fish from the Rio Xingu floodplains, American Society of Ichthyologists and Herpetologists, Wash- Brazilian Amazon. aqua Journal of Ichthyology and ington, DC. Aquatic Biology, 7: 119-122. WOURMS, J. P. 1972. The developmental biology of annual COSTA, W. J. E. M. & BRASIL, G. C. 2008. Simpsonichthys fishes III. Pre-embryonic and embryonic diapause of inaequipinnatus, a new seasonal Killifish from the Tocan- variable duration in the eggs of annual fishes. Journal of tins River basin, Brazil (Cyprinodontiformes, Rivulidae). Experimental Zoology, 182: 389-414. Ichthyological Exploration of Freshwaters, 3: 245-248.

aqua vol. 19 no. 3 - 19 July 2013 122 aqua, International Journal of Ichthyology

Hemiscyllium halmahera, a new species of Bamboo Shark (Hemiscylliidae) from Indonesia

Gerald R. Allen1, Mark V. Erdmann2, 4 and Christine L. Dudgeon3

1) Department of Aquatic Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, Perth, Western Australia 6986. Email: [email protected] 2) Conservation International Marine Program, Jl. Dr. Muwardi No. 17, Renon, Denpasar 80235, Bali, Indonesia 3) University of Queensland, School of Veterinary Sciences, Gatton, Queensland, 4343, Australia 4) California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118, USA

Received: 28 April 2013 – Accepted: 21 June 2013

Abstract großen dunklen U-förmigen Fleck mit mehr oder weniger Hemiscyllium halmahera new species is described from durchgehendem weißen Rand im unteren Teil und einer two specimens, 656-681 mm TL, collected at Ternate, senkrechten Reihe von drei kleineren Gruppen aus 2-3 Halmahera, Indonesia. The new species is clearly differen- dunklen vieleckigen Markierungen besteht. Im allge- tiated on the basis of colour pattern. Its features include a meinen Erscheinungsbild ähnelt die neue Art stark H. galei general brown colouration with numerous clusters of von der Cenderawasih-Bucht, West-Papua, deren Vertreter mainly 2-3 dark polygonal spots, widely scattered white sich aber durch 7-8 große, waagerecht längliche dunkle spots in the matrix between dark clusters, relatively few (< Flecken an der Unterseite zwischen Abdomen und Basis 10), large dark spots on the interorbital/snout region, a der Schwanzflosse unterscheiden, sowie durch eine pair of large dark marks on the ventral surface of the head, Gruppe einheitlich dunkler Flecken hinter dem Kopf und and a fragmented post-cephalic mark consisting of a large gewöhnlich 25 dunkle Flecken auf der Kopfoberseite. U-shaped dark spot with a more or less continuous white margin on the lower half, followed by a vertical row of Résumé three, smaller clusters of 2-3 polygonal dark marks. The Hemiscyllium halmahera, une nouvelle espèce, est décrit sur new species is most similar in general appearance to H. base de deux spécimens, 656-681 mm de LT, collectés à Ter- galei from Cenderawasih Bay, West Papua, which differs in nate, Halmahera, Indonésie. La nouvelle espèce se distingue having 7-8 large, horizontally elongate dark spots on the clairement par le patron de coloration. Ses caractéristiques lower side between the abdomen and caudal-fin base, a comprennent une coloration d’ensemble brune avec nombre cluster of solid dark post-cephalic spots, and usually about de groupes de généralement 2-3 taches polygonales som- 25 dark spots on the upper surface of the head. bres, des taches blanches largement disséminées sur la matrice parmi des groupes sombres, relativement peu Zusammenfassung (moins de 10) de grandes taches foncées dans la région in te - Beschrieben wird die neue Art Hemiscyllium halmahera rorbitale/rostrale, une paire de grandes marques sombres sur auf der Grundlage von zwei Exemplaren mit 656-681 mm la partie ventrale de la tête et une marque fragmentée post - TL (Gesamtlänge), die bei Ternate, Halmahera, in Indone- céphalique qui consiste en une grande tache sombre en for- sien gefangen wurden. Sie sind durch ihr Farbmuster ein- me de U avec une marge blanche plus ou moins continue deutig unterscheidbar. Zu den Merkmalen der neuen Art sur la moitié inférieure, suivie d’une rangée verticale de trois gehören eine allgemein bräunliche Farbgebung mit zahlre- amas plus petits de 2-3 marques polygonales foncées. En ap - ichen Gruppen von meist 2-3 dunklen vieleckigen parence générale, la nouvelle espèce évoque le plus H. galei Flecken, weit verteilte weiße Flecken in der Grundfarbe de Cenderawasih Bay, West Papua, qui diffère par la pré - zwischen den schwarzen Fleckengruppen, relativ wenige (< sence de 7-8 grandes taches sombres étalées à l’horizontale 10) große dunkle Flecken in der Zwischen-Augen- und sur le côté inférieur, entre l’abdomen et la base de la caudale, Schnauzenregion, ein Paar großer dunkler Zeichen an der par un ensemble de taches postcéphaliques foncées et, gé - bauchwärtigen Oberfläche des Kopfes sowie ein zusam- néralement, d’environ 25 taches sombres sur la partie mengesetztes Zeichen hinter dem Kopf, das aus einem supérieure de la tête.

123 aqua vol. 19 no. 3 - 19 July 2013 Hemiscyllium halmahera, a new species of Bamboo Shark (Hemiscylliidae) from Indonesia

Sommario ing” gait while foraging for benthic invertebrates Hemiscyllium halmahera nuova specie è descritta sulla and fishes. Although biological data is sparse for base di due esemplari, di 656-681 mm TL, raccolti a Ter- most species, they are generally oviparous with nate, Halmahera, Indonesia. La nuova specie è chiara- elliptical egg capsules being deposited on the reef. mente distinguibile sulla base della colorazione. Questa Hatchlings are rarely encountered, but the few include una tonalità bruna diffusa con numerosi grappoli di 2-3 macchie poligonali scure, macchie bianche spar - reported specimens were about 15 cm TL (Com- pagliate nella matrice tra i grappoli scuri, relativamente pagno, 2001). poche (<10) grandi macchie scure sulla regione interor- Due to their reproductive mode, limited swim- bitale/muso, un paio di grandi screziature scure sulla ming ability, and poor dispersal capability most superficie ventrale della testa e una in sede post-cefalica, species have restricted distributions. The two Aus- frammentata, composta da una grande macchia scura a tralian species, H. ocellatum and H. trispeculare, are forma di U con un margine bianco più o meno continuo the most widespread, ranging around most of the nella metà inferiore, seguita da una fila verticale di tre northern half of the continent. The remaining grappoli minori di 2-3 macchie poligonali scure. La nuova specie è molto simile nell’aspetto generale a H. galei della species, with the exception of the new species Cenderawasih Bay, Papua occidentale, che si distingue per described below, have restricted regional New avere 7-8 grandi macchie scure, allungate orizzontalmente Guinea distributions (indicated in parentheses; sul lato inferiore tra l'addome e la base della pinna caudale, also see Fig. 1): H. freycineti (Raja Ampat Islands, un gruppo di ben definite macchie scure post-cefaliche e di and possibly adjacent West Papua mainland), H. solito per circa 25 macchie scure sulla superficie superiore galei (Cenderawasih Bay, West Papua), H. hall- della testa. stromi (Gulf of Papua to Milne Bay Province, Papua New Guinea), H. henryi (Kaimana coastline INTRODUCTION and Triton Bay, West Papua), H michaeli (Milne The family Hemiscylliidae, commonly known as Bay and Oro provinces, Papua New Guinea), and bamboo, epaulette, or walking sharks, contains H. strahani (Madang along north coast of Papua small, slender sharks characterised by nasoral and New Guinea to Jayapura, Papua Province, Indone- perioral grooves, short barbels, a small transverse sia). All of these species were illustrated and dis- mouth below the eyes, two similar-sized dorsal fins, cussed by Allen & Erdmann (2012). and a long slender tail. Only two genera are known, The current paper describes a new species from both described by Müller & Henle (1837 & 1838), Halmahera, the first record for the genus from Chiloscyllium with seven Indo-Pacific species, and beyond the Australia-New Guinea region. Two Hemiscyllium with nine species, including the new specimens and tissue samples of each were col- taxon described herein. The family was last lected by M. Erdmann during a night dive at the reviewed by Compagno (2002), who recognized island of Ternate in the Halmahera group of five Hemiscyllium: H. freycineti (Quoy & Gaimard, islands, Indonesia. Subsequent DNA analysis indi- 1824), H. hallstromi Whitley, 1967, H. ocellatum cates it is a separate species. (Bonnaterre, 1788), H. strahani Whitley, 1967, and H. trispeculare Richardson, 1843. Two additional MATERIALS AND METHODS species, H. galei and H. henryi were described by Technical terms and measurements mainly follow Allen & Erdmann (2008) from the Bird’s Head those explained and illustrated by Compagno Peninsula of western New Guinea (West Papua (2001). Snout length is the distance from the snout Province, Indonesia), and a third addition, H. tip to the anterior edge of the mouth. Subcaudal michaeli was described by Allen & Dudgeon (2010) length is the combined measurement of the upper from eastern Papua New Guinea. The members of postventral caudal-fin margin and terminal caudal- the genus are morphologically very similar, and fin margin. Total length and head length are abbre- unfortunately most species are poorly represented viated as TL and HL respectively. Vertebral counts in museum collections, precluding detailed mor- were obtained from radiographs. Vertebral counts phological/meristic comparisons. Therefore, colour and measurements for the holotype are given first patterns and genetic differences remain the most followed by the range for paratypes (in parenthe- valuable “tools” for distinguishing species. ses) in the description below. Type specimens are Sharks of the genus Hemiscyllium are small (usu- deposited at Museum Zoologicum Bogoriense, ally under about 80 cm), nocturnally active, bot- Cibinong, Indonesia (MZB) and the Western Aus- tom-living animals, which exhibit a peculiar “walk- tralian Museum, Perth (WAM). aqua vol. 19 no. 3 - 19 July 2013 124 Gerald R. Allen, Mark V. Erdmann and Christine L. Dudgeon

Tissue samples were obtained for both type spec- 7 min. The PCR products were cleaned using Iso- imens of the new taxon from Ternate, Indonesia. late PCR and Gel Kit (Bioline). Sequences were This was compared with samples of H. freycineti, conducted in both forward and reverse directions H. galei, H. henryi, H. michaeli, and H. ocellatum, using Big Dye Terminator v3.1 Cycle Sequencing which had been previously analysed (Allen and Kit (Applied Biosystems) following manufacturer’s Erdmann 2008; Allen and Dudgeon, 2010). Total instructions. Sequencing products (20ul) were pre- genomic DNA was extracted from 25 mg of fin tis- cipitated by adding 5 µl EDTA (125mM, pH8.0) sue using the DNeasy Tissue Extraction Kit (Qia- and 60 µl 100% ethanol and centrifuged for 45 gen) following the instructions of the supplier. A min at 3000 rpm (30 cm diameter rotor). The pel- fragment including partial mitochondrial NADH let was washed with 60 µl 70% ethanol and cen- Dehydrogenase subunit 4 gene (ND4), tRNA-His trifuged for 15 min at 3000 rpm and then products and tRNA-Ser genes was amplified through poly- resolved on a 3730xl Genetic Analyser (Applied merase chain reaction (PCR) using the primers: Biosystems). ND4-F: 5’ - CACCTATGACTACCAAAAGCT- Sequences were aligned using Sequencher version CATGTAGAAGC - 3’ (Arevalo et al. 1994) and 4.6 (Gene Codes). All molecular statistical analyses H12293-Leu-R: 5’ – TTGCACCAAGAG- were conducted using Mega version 5 (Tamura et TTTTT GGTTCCTAAGACC - 3’ (Inoue et al. al. 2011). Maximum likelihood analysis was used 2001). Reactions were conducted in 30 µl total to assess the model of best fit for the nucleotide amounts and consisted of: 10 µM each primer, 400 substitution. Both Bayesian Information Criterion µM of each dNTP, 3 units Taq polymerase, 1 x (BIC) and Akaike’s Information Criterion cor- PCR Buffer (Qiagen) and 30-50 ng extracted rected for small sample sizes (AICc) ranked the DNA. PCRs were conducted on 9700 Perkin Tamura-Nei substitution model (Tamura & Nei Elmer thermocyclers and consisted of an initial 1993) with a discrete Gamma distribution denaturation step at 95°C for 5 min, followed by (TN93+G; G=0.38) as having the best fit to the 30 cycles of 95°C for 15 sec, 56°C for 30 sec and data. The Tamura-Nei (Tamura & Nei 1993) sub- 72°C for 1 min, and a final extension at 72°C for stitution model accounts for variable base frequen-

Fig. 1. Map of New Guinea and Halmahera showing collection/observation locations of resident species of Hemiscyllium: H. freycineti (yellow circles), H. galei (white star), H. henryi (yellow star), H. hallstromi (white squares), H. halmahera (green circles), H. strahani (red squares), and H. michaeli (red circles).

125 aqua vol. 19 no. 3 - 19 July 2013 Hemiscyllium halmahera, a new species of Bamboo Shark (Hemiscylliidae) from Indonesia cies, transition rates and rate variation among sites. congeners by a unique colour pattern, particularly The TN93+G model was used to construct phylo- a light brown background colour with numerous genetic trees with maximum likelihood analysis. clusters of mainly 2-3 dark polygonal spots, widely Maximum parsimony trees were also constructed scattered white spots in the matrix between dark with a specimen of Chiloscyllium punctatum as the clusters, relatively few (< 10), large dark spots on outgroup taxon. Confidence in tree topology was the interorbital/snout region, pair of large dark evaluated by bootstrapping across 1000 bootstrap marks on ventral surface of head, and fragmented replicates (Felsenstein 1985). Pairwise distances post-cephalic mark consisting of a large U-shaped within and between putative taxa were calculated dark spot with more or less continuous white mar- using the TN93+G model with 1000 bootstrap gin on lower half, followed posteriorly by vertical replicates. row of three, smaller clusters of 2-3 polygonal dark marks. It lacks diagnostic colour pattern features that typify the other known species in the genus, Hemiscyllium halmahera n. sp. including a black hood or face-mask marking on Halmahera Epaulette Shark the head (H. strahani), large intensely black ovate (Figs 2-7; Tables I-II) spots on body (H. hallstromi), ocellated shoulder spot and numerous small black spots on body (H. Holotype: MZB 21248, male, 681 mm TL, north- ocellatum), ocellated shoulder spot with two curved western Ternate, 00°50.958’N, 127°18.717’E, black marks on posterior edge and numerous dif- Halmahera, Indonesia, 10 m, captured by hand, fuse dark spots on head body and fins (H. trispecu- M. Erdmann, 1 May 2012. lare), row of 7-8 large, horizontally ovate dark spots Paratype: WAM P. 33784-001, female, 656 mm on lower side between abdomen and caudal-fin TL, collected with holotype. base (H. galei), “double-ocellus” shoulder marking Diagnosis: A species of bamboo shark belonging (H. henryi), diffuse shoulder marking, numerous to the genus Hemiscyllium, distinguished from all dark spots on dorsal surface of head, and pair of

Fig. 2. Underwater photograph of Hemiscyllium halmahera, male holotype, Ternate Island, Halmahera, Indonesia at depth of 10 m. Photo by M. V. Erdmann. aqua vol. 19 no. 3 - 19 July 2013 126 Gerald R. Allen, Mark V. Erdmann and Christine L. Dudgeon closely spaced dark post-cephalic ocelli surrounded orly; precaudal length 1.3 and HL 6.7 (7.5-8.0) in by white halo (H. freycineti), and dense pattern of TL; head height (at pectoral-fin origin) 1.2 in leopard-like dark spots (H. michaeli). greatest width of head; eye length 3.4 (3.2) in Description: Total vertebral centra 195 (includes snout length, eye height 1.9 (2.7) in eye length; 39-40 monospondylous centra, 101-102 diplos - fleshy interorbital space 1.4 and bony interorbital pondylous precaudal centra, and 54 caudal centra); space 1.7 in snout length; snout blunt and short, body and tail relatively slender, tapering posteri- snout tip to eye 2.2 (2.4), snout tip to mouth 4.6

Fig. 3. Underwater photographs of Hemiscyllium halmahera: upper – approximately 650 mm TL, Bacan Island, Halmahera, Indonesia at depth of 10 m. Photo by J. Yonover. Lower – approximately 700 mm TL, Weda Bay, Halmahera, at depth of 5 m. Photo by T. Mulder.

127 aqua vol. 19 no. 3 - 19 July 2013 Hemiscyllium halmahera, a new species of Bamboo Shark (Hemiscylliidae) from Indonesia

Table I. Proportional measurements (as percentage of total orly, the first 3.9 and fifth 2.7 (2.8) in snout length) for type specimens of Hemiscyllium halmahera. length. Mouth small and transverse, positioned well for- Holotype Paratype ward on ventral surface of head, its width 1.2 (1.3) Measurement MZB WAM in snout length; short barbel on each side of ven- 21248 P. 33784 tral snout, its length 5.2 (4.6) in snout length; maximum width of lower labial flap 5.0 (5.3), Sex male female Total length (mm) 681 656 length of postoral fold (upper labial furrow) 3.4 Precaudal length 78.4 76.5 (3.1), length of lower labial furrow 4.5 (5.3), all in Head width 8.8 8.3 snout length; teeth pavement-like, composed of Head depth 7.3 6.7 numerous rows; individual teeth broad-based with Preanal body depth 5.3 5.2 single posteriorly-directed cusp, the cusps of inner- Snout - pectoral-fin origin (HL) 12.5 13.4 most rows more developed. Snout - 1st gill slit 11.5 11.9 Pre-first dorsal length 2.6 (2.5), prepelvic length 1st to 5th gill slit 5.0 4.9 3.4 (3.2), snout to vent length 3.1 (3.0), vent to First gill slit height 1.5 1.4 anal-fin origin 2.3 (2.4), vent to caudal-fin length Fifth gill slit height 2.1 2.0 Eye diameter (horizontal) 1.7 1.8 1.3, all in TL. Pectoral fins below gill openings, Eye diameter (vertical) 0.9 0.7 their length 1.1 (1.2) in HL; pelvic fins immedi- Bony interorbital width 3.5 3.3 ately anterior to vertical line passing through first Fleshy interorbital width 4.1 3.9 dorsal-fin origin, their length 1.2 (1.4) in HL; dor- Snout to eye (snout length) 5.8 5.7 sal fins positioned well back on body, first and sec- Snout to spiracle 6.8 6.6 ond dorsal fins nearly equal in height; first dorsal- Snout to mouth 2.7 2.3 fin base 1.5 (1.8) in HL, first dorsal-fin height 1.1 Lower labial furrow length 1.3 1.1 in first dorsal-fin base; free margin of first dorsal Maximum width lower labial flap 1.1 1.1 fin 2.1 (1.4) in first dorsal-fin height; interdorsal Postoral fold 1.7 1.8 space 1.1 in HL; second dorsal-fin base 1.6 (1.7) in Mouth width 4.9 4.2 Barbel length 1.1 1.2 HL; second dorsal-fin height 1.0 (1.1) in second Snout -1st dorsal origin 37.9 40.2 dorsal-fin base; free margin of second dorsal fin 2.0 Snout - pelvic origin 29.0 31.2 (1.9) in second dorsal-fin height; long and low anal Snout - anal opening 32.2 33.6 fin just anterior to caudal fin; anal-fin base 1.4 Anal opening - anal-fin origin 42.9 40.8 (1.7) in HL, anal-fin height 3.3 (2.7) in anal-fin Anal opening - tail tip 68.8 67.1 base; free margin of anal fin 1.5 (1.8) in anal-fin Interdorsal distance 11.3 11.8 height; elongate and thick precaudal tail (section of Pectoral-fin length 11.7 11.0 body between anus and caudal fin), its depth at Pelvic-fin length 10.4 9.9 level of anal-fin origin 2.2 (2.6) in head length; 1WAM dorsal fin base 8.5 7.4 1st dorsal-fin height 8.1 6.5 subcaudal length 6.2 (5.7) in TL. 1st dorsal-fin free margin 4.9 4.5 Clasper of adult male holotype (Fig. 4A) relatively 2nd dorsal-fin base 8.1 7.7 stout and elongate, its length greater than first dor- 2nd dorsal-fin height 7.4 7.4 sal-fin height. 2nd dorsal-fin free margin 3.7 4.0 Colour in life (Figs. 2-3): generally brown Anal-fin base 9.2 7.8 on snout, dorsally on head and body, grading to Anal-fin height 2.8 2.9 golden brown on lower side of body, and white on Anal-fin free margin 1.9 1.6 ventral surface; numerous clusters of mainly 2-3 Subcaudal 16.2 17.6 dark polygonal spots (largest about equal to eye) Clasper length (inner) 8.9 - Clasper length (outer) 6.3 - and widely scattered, much smaller, white spots in brownish matrix between dark clusters; two (paratype) or three (holotype), progressively (5.8), snout tip to spiracle 1.8 (2.0), snout tip to smaller dark grey brown saddles across back from first gill slit 1.1, all in HL; gill slits on rear part of rear edge of head to dorsal-fin base, a similar sad- head, above to slightly anterior of pectoral-fin base; dle between dorsal fins, and 4-5 additional dark distance between first and fifth gill slit 2.5 (2.7) in saddles on dorsal edge of tail; four large saddles on HL; height of gill slits gradually increasing posteri- body with narrow white or pale grey anterior and aqua vol. 19 no. 3 - 19 July 2013 128 Gerald R. Allen, Mark V. Erdmann and Christine L. Dudgeon

Table II. Inter-specific pairwise distance matrix (below diagonal) calculated from the TN93+G model with corresponding matrix (above diagonal) of SE estimates (bootstrap method, 1000 replicates).

H. halmahera H. michaeli H. freycineti H. galei H. henryi H. ocellatum C. punctatum

H. halmahera - 0.0052 0.0058 0.0066 0.0059 0.0065 0.0219 H. michaeli 0.0201 - 0.0056 0.0060 0.0057 0.0068 0.0237 H. freycineti 0.0229 0.0226 - 0.0043 0.0036 0.0072 0.0202 H. galei 0.0286 0.0254 0.0130 - 0.0047 0.0077 0.0209 H. henryi 0.0229 0.0227 0.0103 0.0184 - 0.0073 0.0203 H. ocellatum 0.0314 0.0329 0.0372 0.0401 0.0371 - 0.0215 C. punctatum 0.1627 0.1747 0.1545 0.1588 0.1547 0.1572 - posterior margins; fragmented post-cephalic mark few (< 10) large (smaller than eye, Fig. 4D), dark (Fig. 4B), consisting of a large U-shaped dark spot spots on interorbital/snout region; each dorsal fin with more or less continuous white margin on with few poorly defined brown spots and pair of lower half, followed by vertical row of three, prominent blackish saddles on anterior edge, the smaller clusters of 2-3 polygonal dark marks; pair lowermost ocellus-like with white margin around of large round to oval dark marks (Fig. 4C) on ven- lower edge; pectoral and pelvic fins with 8-10 and tral surface of head at about level of spiracle; lower 6-11 variable-sized brown spots respectively on side of head, below and slightly posterior to spira- dorsal surface and narrow white posterior margin. cle with 2-3 large, irregular dark spots; relatively Colour in alcohol (Figs 5-7) similar to the

Fig. 4A-D. Diagnostic features of Hemiscyllium halmahera: A. claspers of male holotype; B. post-cephalic marking of holotype; C. ventral surface of head of female paratype; D. dorsal surface of head of male holotype. Photos by G. R. Allen.

129 aqua vol. 19 no. 3 - 19 July 2013 Hemiscyllium halmahera, a new species of Bamboo Shark (Hemiscylliidae) from Indonesia

Fig. 5. Hemiscyllium halmahera, preserved male holotype, 681 mm TL, Ternate, Halmahera, Indonesia. Photo by G. R. Allen.

Fig. 6. Hemiscyllium halmahera, dorsal view of preserved holotype, 681 mm TL, Ternate, Halmahera, Indonesia. Photo by G. R. Allen.

Fig. 7. Hemiscyllium halmahera, preserved female paratype, 656 mm TL, Ternate, Halmahera, Indonesia. Photo by G. R. Allen. aqua vol. 19 no. 3 - 19 July 2013 130 Gerald R. Allen, Mark V. Erdmann and Christine L. Dudgeon

Fig. 8. Underwater photograph of Hemiscyllium michaeli, approximately 600 mm TL, Milne Bay Province, Papua New Guinea. Photo by B. Halstead.

Fig. 9. Underwater photograph of Hemiscyllium freycineti, approximately 600 mm TL, Kri Island, Raja Ampat Islands, West Papua Province, Indonesia. Photo by G. R. Allen.

131 aqua vol. 19 no. 3 - 19 July 2013 Hemiscyllium halmahera, a new species of Bamboo Shark (Hemiscylliidae) from Indonesia live colour provided above, except the ground differences for the other members of the genus. colour is tan to reddish brown and the dark spot- Pairwise distances between the Hemiscyllium ting on the head, body, and fins is less intense. species and the outgroup Chiloscyllium punctatum DNA Analysis: In total, we analysed a 792 base (dave= 0.1559) were over an order of magnitude pair (bp) alignment of the mitochondrial ND4 higher than some of the intra-genus distances. gene from 18 Hemiscyllium individuals and one Only H. ocellatum showed intra-species distance Chiloscyllium punctatum individual. There were (d = 0.0013) which was an order of magnitude two new samples for H. halmahera that were com- lower than the intra genus distances. pared with the 16 sequences included in the previ- The maximum likelihood (ML) and maximum ous genetic analyses from Allen and Erdmann parsimony (MP) trees both clearly separated out (2008) (H. henryi = 4; H. ocellatum = 5; H. H. halmahera from the morphologically similar H. freycineti = 4; H. galei = 2) and Allen and Dudgeon galei. H. halmahera was positioned in a basal posi- (2010) (H. michaeli = 1). The two H. halmahera tion of the Hemiscyllium samples with H. michaeli. samples had identical sequences. A total of 6 hap- The phylogenetic analyses produced identical trees lotypes were detected for all Hemiscyllium species for most of taxa, but could not resolve the relative with all species having one haplotype, except for H. positioning of H. halmahera and H. michaeli with ocellatum with two haplotypes. In the total Hemi - ML analyses placing H. michaeli in the most basal scyllium alignment there were 50 variable sites and position (with low bootstrap support = 49%) and 43 parsimony-informative characters. Nucleotide MP analyses reversing this relationship (bootstrap frequencies of H. halmahera were similar to the support = 62%; Fig. 11). other species and average frequencies for the com- Distribution and habitat: The new species is bined Hemiscyllium samples were as follows: A = known with certainty from satellite islands off the 32, C = 24, G = 0.11, T = 32. Pairwise distances west coast of Halmahera, Indonesia and in Weda (d) between the Hemiscyllium species ranged from Bay in South Halmahera. The type specimens were 0.0103-0.0401. The H. halmahera samples clearly collected at night in 10 m depth, nestling under differed from all the other Hemiscyllium species coral heads that were sparsely scattered on a steep, with pairwise distances ranging from 0.0201- black volcanic sand slope off northwestern Ternate 0.0314 which fall within the range of inter-species Island. It has also been photographed off south-

Fig.10. Underwater photograph of Hemiscyllium galei, approximately 600 mm TL, Cenderawasih Bay, West Papua Province, Indonesia. Photo by G. R. Allen. aqua vol. 19 no. 3 - 19 July 2013 132 Gerald R. Allen, Mark V. Erdmann and Christine L. Dudgeon western Halmahera at Proco Island (00°25.264’S, Hemiscyllium beyond the Australia/New Guinea 127°44.264’E), which lies in the Proco Strait region. Although Halmahera lies in close proxim- between Bacan and mainland Halmahera; at Bacan ity to New Guinea, the current study extends the Island (00°20.371’S, 127°18.153’E); and in Weda known boundary for the genus approximately 300 Bay in southern Halmahera (00°27.935’N, 127° km further west. As mentioned previously, Hemi - 56.753’E) . scyllium species appear to have limited dispersal Etymology: The species is named H. halmahera capabilities, including an apparent inability to with reference to the type locality. The name is cross significant deep-water barriers. For example, treated as a noun in apposition. the genus is unknown from the island of New Remarks: The new species is the first record for Britain, which lies only about 80 km from the

Fig. 11. Maximum likelihood (ML) tree (TN93+G) of the ND4 data for six Hemiscyllium species. Bootstrap support for 1000 replicates are shown for ML and Maximum Parsimony (MP) trees respectively. ML and MP trees are identical except for the relative positions of H. halmahera and H. michaeli, which are reversed in the MP phylogeny with bootstrap support of 62% (indicated by *). The outgroup is Chiloscyllium punctatum Müller & Henle, 1838, a member of the other genus in the family Hemiscylliidae.

133 aqua vol. 19 no. 3 - 19 July 2013 Hemiscyllium halmahera, a new species of Bamboo Shark (Hemiscylliidae) from Indonesia

Papua New Guinea mainland. Based on paleogeo- At present, the genus is poorly represented in graphic reconstructions presented by Hall (2002) museum collections and therefore morphometric and Hill & Hall (2003) it seems likely that the data is of limited value in separating the various ancestral population may have colonized Halma- species. However, the new species appears to have hera sometime after 25 MYA when its island-arc an unusually thick tail base (preanal body depth) fragments precursor came within close proximity compared to its congeners. For example, the pre- of the northern New Guinea mainland. Then anal depth for H. halmahera is 2.3-2.6 in the head according to these reconstructions, the component length compared with values of 3.5-4.5 for H. parts of Halmahera underwent a long, slow west- galei, H. henryi, and H. michaeli. However, more ward drift during the Miocene and Pliocene, arriv- specimens are required for most species to fully ing at their present position over the past few mil- evaluate these differences. lion years. Therefore, it seems highly likely that the The species of Hemiscyllium are distinguished ancestral population of H. halmahera was rafted mainly on the basis of colour pattern. The follow- from a distant colonization point to its current ing key will serve to identify the nine known location. Similar scenarios involving dispersal of species. New Guinea freshwater Heteroptera (water strid- ers) via island arc fragments were discussed by Pol- Key to the Species of Hemiscyllium hemus & Polhemus (1998), and may also explain the presence of the bird-of-paradise Semioptera 1a. Head and snout with an abrupt black hood; wallacii on Halmahera, despite the family Paradis- body covered with conspicuous large white aeidae otherwise being considered endemic to the spots (northern New Guinea) ...... island of New Guinea and Australia (Irestedt et al., ...... H. strahani 2009). We moreover note that there are a number 1b. Head and snout light in colour, without a black of examples of restricted range New Guinea reef hood but with conspicuous black spots above fish species whose ranges just penetrate into pectoral fins; body with inconspicuous light Halmahera, including Pseudochromis ammeri Gill, spots or spots absent ...... 2 Allen and Erdmann, 2012, P. matahari Gill, Erd- 2a. Dark spot behind gills relatively small, forming mann and Allen, 2009, P. pylei Randall and a weak ocellus (or semi-ocellate dark spot) and McCosker, 1989, and Pentapodus numberii Allen followed posteriorly by dark brown bar (con- and Erdmann, 2009, as well as a blenny (Ecsenius sisting of 2-3 merged spots in vertical row or 2- randalli Springer, 1991) considered endemic to 3 clusters of fragmented spots)...... 3 Halmahera that is found occasionally in Raja 2b. Dark spot behind gills large, forming 1-2 con- Ampat, West Papua. spicuous white-rimmed ocelli or merged dou- The new species is most similar in colour pattern ble ocelli without dark brown bar immediately to H. galei (Fig. 10) from Cenderawasih Bay, West behind ...... 5 Papua. Both species have characteristic broad dark 3a. Lower side with conspicuous horizontal row of saddles with narrow, white anterior and posterior row of 7-8 large, horizontally-oval, dark spots margins on the dorsal surface of the body, as well (Cenderawasih Bay, West Papua) ...... H. galei as similar post-cephalic markings. However, there 3b. Lower side without row of large, horizontally- are significant differences between the two species. oval, dark spots along lower side ...... 4 Hemiscyllium galei differs in having a row of 7-8 4a. Ventral surface of head with pair of large dark large, horizontally ovate spots on the lower side spots; snout region of adult with only a few between the abdomen and caudal-fin base and has small dark spots (Halmahera)...... more spots (usually about 25) on the dorsal surface ...... H. halmahera of the head. In contrast, H. halmahera differs in 4b. Ventral surface of head without pair of large having far fewer head spots (usually less than 10) dark spots; snout region of adult with numer- and possesses a pair of large dark spots on the ven- ous small dark spots (Raja Ampat Islands, West tral surface of the head. Although the general con- Papua) ...... H. freycineti figuration of the post-cephalic spot is similar, that 5a. Head, body, and fins covered with polygonal, of H. halmahera consists of clusters of fragmented leopard-like spots (eastern Papua New Guinea) spots (with a U-shaped main spot) compared to ...... H. michaeli the solid spots of H. galei. 5b. Head body, and fins covered with numerous aqua vol. 19 no. 3 - 19 July 2013 134 Gerald R. Allen, Mark V. Erdmann and Christine L. Dudgeon

spots, but more or less round in shape and not conservation programs to protect this unique polygonal and leopard like ...... 6 species on the reefs of Halmahera. 6a. Body covered with numerous, densely clustered, dark small and large spots that form a reticular network of light base colour between REFERENCES them; dark cross-bands well defined on ventral ALLEN, G. R. & DUDGEON, C. L. 2010. Hemiscyllium surface of tail (northern Australia)...... michaeli, a new species of bamboo shark (Hemiscyllidae) from Papua New Guinea. aqua, International Journal of ...... H. trispeculare Ichthyology 16 (1): 19-30. 6b. Body with fewer large spots that do not form a ALLEN, G. R. & ERDMANN, M. V. 2008. Two new species reticular network of light ground colour of bamboo sharks (Orectolobiformes: Hemiscyllidae) between them; dark cross-bands on tail rela- from western New Guinea. aqua, International Journal of tively weak or not reaching ventral surface...... Ichthyology 13 (3-4): 93-108...... 7 ALLEN, G. R. & ERDMANN, M. V. 2012. Reef Fishes of the 7a. Lateral ocellus surrounded by large black spots; East Indies. Volume 1. Tropical Reef Research, Perth, spots absent on head in front and below eyes Australia, 424 pp. AREVALO, E., DAVIS, S. K. & SITES, J. W. 1994. Mito- (south-eastern Papua New Guinea)...... chondrial-DNA sequence divergence and phylogenetic- ...... H. hallstromi relationships among eight chromosome races of the Scelo- 7b. Lateral ocellus surrounded by relatively small porus-Grammicus complex (Phrynosomatide) in central spots; spots present on head in front and below Mexico. Systematic Biology 43: 387-418. eyes ...... 8 BONNATERRE, J. P. 1788. Tableau encyclopédique et method- 8a. Lateral ocellus composed of single large, round ique des trois règnes de la nature... Ichthyologie. Panck- spot surrounded by distinct white halo; dark oucke, Paris. spot absent at origin of pectoral and pelvic fins; COMPAGNO, L. J. V. 2001. Sharks of the world. An anno- tated and illustrated catalogue of shark species known to numerous small dark spots usually present on date. Volume 2. Bullhead, mackerel and carpet sharks (Het- dorsal fins (northern Australia) ...... erodontiformes, Lamniformes and Orectolobiformes). FAO ...... H. ocellatum Species Catalogue for Fishery Purposes. No. 1, Vol. 2. 8b. Lateral ocellus usually composed of double, FAO, Rome, 269 pp. merged ocelli surrounded by poorly defined FELSENSTEIN, J. 1985. Confidence limits on phylogenies: white halo; dark spot present at origin of pec- an approach using the bootstrap. Evolution 39: 783-791. toral and pelvic fins; dorsal fins usually without HALL, R. 2003. Cenozoic geological and plate tectonic small dark spots or if present they are restricted evolution of SE Asia and the SW Pacific: computer-based reconstructions, model, and animations. Journal of Asian to basal portion of fins (Triton Bay region, Earth Sciences 20: 353-434. West Papua)...... H. henryi HILL, K. C. & HALL, R. 2003. Mesozoic-Cenozoic evolution of Australia’s New Guinea margin in a west Pacific ACKNOWLEDGEMENTS context. In Hillis R.R. & Müller, R.D. (eds.) Evolution We thank Graham Abbott for first calling our and Dynamics of the Australian Plate, pp. 265-290. Geo- attention to the presence of Hemiscyllium on logical Society of Australia Special Publication 22 and Halmahera with his photo from Proco Strait, and Geological Society of America Special Paper 372. Alberth Reija, Tony Rhodes and the outstanding INOUE, J. G., MIYA, M., TSUKAMOTO, K. & NISHIDA, M. 2001. A mitogenomic perspective on the basal teleostean crew of the Damai II live-aboard for their dive sup- phylogeny: Resolving higher-level relationships with port in collecting the type specimens; dive guide longer DNA sequences. Molecular Phylogenetics and Evo- Gusti in particular deserves mention for his sharp lution 20: 275-285. eyes spotting the sharks at night. Thanks are due IRESTEDT, M., JØNSSON, K. A., FJELDS, Å, J., CHRISTIDIS, Dita Cahyani and the Udayana University staff for L. & ERICSON, P. G. 2009. An unexpectedly long history assistance in x-raying the holotype, and Renny of sexual selection in birds-of-paradise. BMC Evolution- Hadiaty of LIPI for assistance in curating the holo- ary Biology 9: 235. type. We also thank Jeff Yonover and Tom Mulder MÜLLER, J. & HENLE, F. G. J. 1837. Ueber die Gattun- gen der Plagiostomen. Archiv für Naturgeschichte 3: for the use of their excellent images of the new 394-401, 434. species, and Rob Sinke for further information on MÜLLER, J. & HENLE, F. G. J. 1838. On the generic char- the species in Weda Bay. Finally, we thank the gov- acters of cartilaginous fishes, with descriptions of new ernment of Maluku Utara for permitting our trip genera. Magazine of Natural History (new series) 2: 33- and for their efforts to develop marine tourism and 37, 88-91.

135 aqua vol. 19 no. 3 - 19 July 2013 Hemiscyllium halmahera, a new species of Bamboo Shark (Hemiscylliidae) from Indonesia

POLHEMUS, D. A. & POLHEMUS, J. T. 1998. Assembling TAMURA, K., PETERSON, D., PETERSON, N., STECHER, G., New Guinea: 40 million years of island arc accretion as NEI, M., KUMAR, S. 2011. MEGA5: Molecular evolu- indicated by the distributions of aquatic Heteroptera tionary genetics analysis using maximum likelihood, evo- (Insecta). In Hall & Holloway (eds.) Biogeography and lutionary distance, and maximum parisomony methods. Geological Evolution of SE Asia, pp. 327-340. Blackhuys Molecular Biology and Evolution 28: 2731-2739. Publishers, Leiden, The Netherlands. TAMURA, K. & NEI, M. 1993. Estimation of the number QUOY, J. R. C. & GAIMARD, J. P. 1824-25. Description des of nucleotide substitutions in the contral region of mito- Poissons. Chapter IX. In: Freycinet, L. de, Voyage autour chondrial DNA in humans and chimpanzees. Molecular du Monde...exécuté sur les corvettes de L. M. “L’Uranie” Biology and Evolution 10: 512-526. et “La Physicienne,” pendant les années 1817, 1818, WHITLEY, G. P. 1967. Sharks of the Australasian region. 1819 et 1820. Paris. Australian Zoologist 14 (2): 173-188. RICHARDSON, J. 1843. Icones piscium, or plates of rare fishes. Part I. Richard and John E. Taylor, London.

aqua vol. 19 no. 3 - 19 July 2013 136 aqua, International Journal of Ichthyology

Melanotaenia sneideri, a New Species of Rainbowfish (Melanotaeniidae), from West Papua Province, Indonesia

Gerald R. Allen1 and Renny K. Hadiaty2

1) Department of Aquatic Zoology, Western Australian Museum, Locked Bag 49, Welshpool DC, Perth, Western Australia 6986. Email: [email protected] 2) Indonesian Institute of Sciences (LIPI), Research Centre for Biology, Division of Zoology, Museum Zoologicum Bogoriense (MZB), Jalan Raya Bogor, Km 46, Cibinong 16911, Indonesia

Received: 13 May 2013 – Accepted: 03 July 2013

Abstract snei deri, est décrite, originaire de la Bomberai Peninsula au A new species of melanotaeniid rainbowfish, Melanotae- sud-ouest de la région du Bird’s Head en Nouvelle-Guinée nia sneideri, is described from the Bomberai Peninsula in occidentale (West Papua Province, Indonésie). Ce nouveau the southwestern Bird’s Head region of western New taxon est décrit sur base de 25 spécimens, de 15,9 à 80,1 mm Guinea (West Papua Province, Indonesia). The new taxon de LS, collectés dans un petit cours d’eau alimenté par une is described on the basis of 25 specimens, 15.9-80.1 mm source karstique, dans un petit bassin lacustre temporaire à SL, collected from a karst spring-fed creek in a small une altitude de 1.050 m, dans les monts Kumawa. Il se di- ephemeral lake basin at an altitude of 1,050 m in the stingue de ses congénères par la combinaison suivante: le cor- Kumawa Mountains. It is distinguished from congeners by ps d’un rouge lumineux, des nageoires dorsale, anale et ven- a combination of the bright red body colour, dark brown trale brun foncé à noires et un corps relativement haut to blackish dorsal, anal, and pelvic fins, and relatively deep (jusqu’à au moins 42,3% de la LS) pour les mâles adultes. Les body (to at least 42.3 % of SL) of adult males. Additional caractéristiques diagnostiques comprennent 18-20 branchio- diagnostic features include 18-20 gill rakers on the first spines sur le premier arc branchial, 15-16 écailles circumpé- branchial arch, 15-16 circumpeduncular scales, and an donculaires et l’absence de dents vomérines ou un groupe de absence of vomerine teeth or a small, inconspicuous patch petites dents vomérines rudimentaires ou imperceptibles. of rudimentary vomerine teeth. Sommario Zusammenfassung Una nuova specie di pesce arcobaleno, Melanotaenia snei- Beschrieben wird eine neue Art der Regenbogenfische, deri, è descritta dalla penisola Bomberai della regione sud- Melanotaenia sneideri, von der Halbinsel Bomberai in der occidentale di Bird’s Head, Nuova Guinea occidentale südwestlichen Vogelkopf-Gegend des westlichen Neuguinea (Provincia di West Papua, Indonesia). Il nuovo taxon è (Provinz Westpapua, Indonesien). Das neue Taxon wird auf descritto sulla base di 25 esemplari di 15.9-80.1 mm SL, der Grundlage von 25 Exemplaren mit 15,9-80,1 mm SL raccolti presso un torrente alimentato da una sorgente car- beschrieben, die in den Kumawa-Bergen in 1.050 m Höhe sica in un piccolo bacino lacustre effimero, a un'altitudine aus einem kleinen zeitweiligen Seebecken in einem quellge- di 1050 metri sulle montagne Kumawa. Si distingue dai speisten Flüsschen in Karstgelände gesammelt wurden. Sie congeneri per una combinazione di caratteri, in particolare lassen sich von anderen Angehörigen der Gattung durch un brillante color rosso del corpo, pinne dorsale, anale e Farbmerkmale unterscheiden: leuchtend roter Rumpf, pelviche dal marrone scuro al nerastro e per il corpo dei dunkelbraune bis schwärzliche Rücken-, After und Bauch- maschi adulti relativamente profondo (almeno al 42,3% flossen, sowie durch einen verhältnismäßig tiefen Körper della SL). Altre caratteristiche diagnostiche includono 18- bei den Männchen (bis mindestens 42,3 % der Standard- 20 branchiospine sul primo arco branchiale, 15-16 scaglie länge SL). Zu den weiteren Bestimmungsmerkmalen circumpeduncolari, e l'assenza di denti vomerini o al più la gehören die Zahl von 18-20 Kiemenreusen am ersten presenza di un piccolo, poco appariscente, raggruppa- Kiemenbogen, 15-16 Schuppen um den Schwanzstiel, ganz mento di rudimentali denti vomerini. fehlende oder nur als kleines, unauffälliges Rudiment vorhandene Gaumenzähne. INTRODUCTION Résumé Rainbowfishes (Melanotaeniidae) are among the Une nouvelle espèce de Melanotaeniidae, Melanotaenia most common inhabitants of fresh waters of the

137 aqua vol. 19 no. 3 - 19 July 2013 Melanotaenia sneideri, a New Species of Rainbowfish (Melanotaeniidae), from West Papua Province, Indonesia

Australia and New Guinea region, both in terms of date the separation of the Vogelkop from north- species and number of individuals. They occur in a western Australia as Early Cretaceous, but the huge variety of lotic and lentic habitats. Allen resulting terrane did not move far (Hill & Hall (1995) presented an overview of the family, recog- 2003), probably remaining slightly southwest of nising 55 species, including 13 from Australia, 40 the main section of New Guinea, and possibly from New Guinea, and three, which are shared. undergoing in-place clockwise rotation of up to Recent taxonomic studies resulted in the descrip- 90° (Hamilton 1979). During the Early Tertiary tion of 25 additional species, including three Chi- the original core of the Bird’s Head was expanded latherina Regan, 1914 (Allen & Renyaan 1996a; by the fusion of the continentally-derived Misool Price 1997; Allen & Unmack 2012), two Glos- terrane to its western margin and the arc-related solepis Weber, 1907 (Allen 2001; Allen & Renyaan Tamrau Terrane to its northern edge (Pigram & 1998), 19 Melanotaenia Gill, 1862 (Allen 1996; Davies 1987). This entire composite unit then Allen 1997; Allen & Hadiaty 2011; Allen & moved eastward, colliding with greater New Renyaan 1996b and 1998; Allen & Unmack 2008; Guinea in the Miocene (approximately 5-6 MYA). Allen et al. 2008; Kadarusman et al. 2010, 2011, The former collision zone is very evident today in and 2012; McGuigan 2001), and a new monotypic the form of the stacked anticlines of continental genus, Pelangia Allen, 1998. In addition, several shelf limestone in the Lengguru Fold Belt of the species that were previously considered as syn- “Bird’s Neck” region. It is among the most rugged onyms or subspecies, including Glossolepis kabia landscapes on the island, containing extensive karst (Herre, 1935), Melanotaenia australis (Castelnau, topography with many lakes, forming reservoirs for 1875), and M. rubrostriatus (Ramsay & Ogilby, independent watersheds. Most of the lakes thus far 1886), were elevated to full specific status (Allen et investigated appear to have subterranean drainage al. 2002; Graf & Ohee 2009). Therefore, 82 and harbour endemic rainbowfishes (Allen & species, including the new taxon described below, Hadiaty 2011; Allen 1998; Allen & Renyaan are currently known for the family. 1996b). More recently, Kadarusman et al. (2012) Unmack et al. (2013) provided a genetic analysis described four species of Melanotaenia (M. arguni, of Melanotaeniidae, which indicates three main M. urisa, M. veoliae, and M. wanona) from karst lineages corresponding with the major geographic springs and associated creeks in the vicinity of regions of New Guinea (northern, southern, and Kaimana (03°39.727’S, 133°45.740’E). western). Most of the 22 species endemic to west- The present paper describes an additional karst- ern New Guinea (Bird’s Head region of West spring associated species from the previously unex- Papua Province, Indonesia) appear to be closely plored and isolated Kumawa Mountains (Fig. 1), allied. The majority of these inhabit the Raja which lie approximately 75 km southwest of the Ampat Islands (e.g. Batanta, Misool, Salawati, and aforementioned Kaimana region. The distinctively Waigeo) or have extremely limited local distribu- coloured new species was discovered by Dr. tions, often restricted to lakes, small creeks, and Richard Sneider and collected for the authors dur- springs in karst areas. Of particular interest is the ing March 2013 by Max Ammer, Obed Holago, relatively narrow “Bird’s Neck” isthmus (Fig. 1) and Richard Sneider during the course of a heli- connecting the Bird’s Head Peninsula with the copter reconnaissance of the Kumawa Mountains. remainder of New Guinea, including the Bomberai Peninsula. This area has a tumultuous geologic past MATERIALS AND METHODS and is among the most rugged landscapes on the Counts and measurements that appear in paren- island, containing extensive karst topography char- theses refer to the range for paratypes if different acterized by subterranean drainage and countless from the holotype. Type specimens are deposited at independent watersheds. Museum Zoologicum Bogoriense, Cibinong, Java, Polhemus (2006) gave a detailed summary of the Indonesia (MZB), National Museum of Natural tectonic history of the New Guinea region. Geolo- History, Washington, D.C. (USNM), and Western gists are in general agreement that the core of the Australian Museum, Perth (WAM). Bird’s Head or Vogelkop Peninsula is a section of The methods of counting and measuring are as the Australian craton that became detached from follows: dorsal and anal rays – the last ray of the the main continental mass sometime in the Meso- anal and second dorsal fins is divided at the base zoic (Hamilton 1979). Pigram & Davies (1987) and counted as a single ray; lateral scales – number aqua vol. 19 no. 3 - 19 July 2013 138 Gerald R. Allen and Renny K. Hadiaty of scales in horizontal row from upper edge of pec- caudal concavity – horizontal distance between ver- toral-fin base to caudal-fin base, excluding the ticals at tips of shortest and longest rays. small scales posterior to the hypural junction; transverse scales – number of scales in vertical row (excluding small truncated scales along base of fins) Melanotaenia sneideri, n. sp. between anal-fin origin and base of first dorsal fin; Kumawa Rainbowfish predorsal scales - number of scales along midline of (Figs 2-4; Tables I-II) nape in front of first dorsal fin; cheek scales – total number of scales covering suborbital and preoper- Holotype: MZB 21375, male, 80.1 mm SL, small culum; standard length (SL) – measured from tip of creek in Kumawa Mountains, 03°55.14’S, upper lip to caudal-fin base; head length (HL) – 133°02.873’E, Bomberai Peninsula, West Papua, measured from tip of upper lip to upper rear edge Indonesia, hand nets, R. Sneider, 27 March 2013. of gill opening; caudal peduncle depth is least depth Paratypes (same data as holotype except collected and caudal peduncle length is measured between by Obed Holago): MZB 21376, 11 specimens, two vertical lines, one passing through base of last 14.1-78.3 mm SL; USNM 409975, 4 specimens, anal ray and the other through caudal-fin base; 36.5-79.8 mm SL; WAM P.33872-001, 5 speci-

Fig. 1. Map of West Papua Province, Indonesia. The type locality of Melanotaenia sneideri is shown by the arrow indicating the Kumawa Mountains.

139 aqua vol. 19 no. 3 - 19 July 2013 Melanotaenia sneideri, a New Species of Rainbowfish (Melanotaeniidae), from West Papua Province, Indonesia mens, 37.5-78.3 mm SL; WAM P.33872-002, 4 2.7 (2.7-3.3) in HL; eye diameter 3.6 (3.2-3.6) in specimens, 15.9-69.6 mm SL. HL; interorbital width 2.5 (2.4-2.8) in HL; depth Diagnosis: A species of melanotaeniid distin- of caudal peduncle 2.3 (2.3-2.8) in HL; length of guished by the following combination of characters: caudal peduncle 1.5 (1.4-1.7) in HL. dorsal rays IV to VI-I,13 or 14 (usually V-I,13); anal Jaws nearly equal, mouth oblique, and premaxilla rays I,23-28 (most frequently 24); pectoral rays 13 with an abrupt bend between anterior horizontal to 15 (usually 14); lateral scales 34-36 (usually 35), portion and lateral part; maxilla ends below about predorsal scales 17-18; cheek scales 14-18 (x– = 15.6); anterior edge of eye; maxillary length 2.6 (2.7-3.3) circumpeduncular scales 15-16; total gill rakers on in HL; lips thin; teeth conical with slightly curved first arch 18-20 (usually 18-19); vomerine teeth tips, those on upper jaw mainly on outer surface of absent or present as inconspicuous rudimentary lips; teeth of upper jaw in about 4-5 irregular rows patch; head length 3.6-4.0 18 (x– = 3.8) in SL; depth anteriorly, reduced to 1-2 rows posteriorly, where of caudal peduncle 2.3-2.8 (x– = 2.5) in HL; maxi- exposed when mouth closed; teeth in lower jaw in mum body depth of male (80.1 mm SL) 42.3 % SL; about 7-8 irregular rows anteriorly, reduced to 2-3 colour in life of adult male generally bright red with rows posteriorly; teeth generally absent on vomer bluish head, mainly dark brown to blackish dorsal, and palate, except small vestigial patch of vomerine anal, and pelvic fins, and pinkish orange caudal fin. teeth on one side of holotype and two paratypes; Description: Dorsal rays V-I,13 (IV to VI-I,13 or roof of mouth and upper surface of tongue covered 14); anal rays I,23 (I,23-28); pectoral rays 13/15 with numerous slender papillae. (13-15); pelvic rays I,5; branched caudal rays 15; Scales of body cycloid with scalloped posterior procurrent caudal rays 6 (5-6); lateral scales 35 margin, relatively large, and arranged in regular (33-36); transverse scales 12; predorsal scales 17 horizontal rows; row of small, truncated scales (17-18); prepelvic scales 18 (17-19); cheek scales along bases of dorsal and anal fins; no scales on 14 (14-18; –x = 15.6); circumpeduncular scales 16 membranous portions of fins except several rows of (15-16); total gill rakers on first arch 19 (18-20). Body depth 2.4 (2.6-3.3) in SL; greatest body depth by sex and size class as follows: male – 35-55 mm SL, 33.7-34.5 % SL (x– = 34.1, N = 2); male – 69.6 mm SL, 37.2 % SL; male – 80.1 mm SL (holotype), 42.3 % SL; females – 33-55 mm SL, 29.2-30.7 % SL (x– = 30.1, N = 4); females – 70-84 mm SL, 30.3-38.2 % SL (x– = 34.2, N = 5); head length 3.7 (3.6-4.0) in SL; greatest width of body 2.7 (1.9-2.7) in greatest body depth; snout length A

B Fig. 2A-B. Live photographs of freshly collected holotype of Melanotaenia sneideri, male, 80.1 mm SL. Photos by R. Sneider. aqua vol. 19 no. 3 - 19 July 2013 140 Gerald R. Allen and Renny K. Hadiaty

Table I. Proportional measurements of selected type specimens of Melanotaenia sneideri expressed as % of the standard length.

Holotype Paratype Paratype Paratype Paratype Paratype Paratype Paratype MZB USNM MZB USNM WAM WAM MZB WAM 21375 409975 21376 409975 P.33872 P.33872 21376 P.33872

Sex male female female female male male female female Standard length (mm) 80.1 79.8 78.3 71.6 69.6 45.8 52.7 41.5 Body depth 42.3 34.2 32.7 30.3 37.2 34.5 29.8 30.6 Body width 15.5 16.2 16.5 15.9 15.2 13.5 15.2 13.0 Head length 27.1 25.6 26.6 25.7 27.0 27.3 28.1 26.3 Snout length 10.0 8.1 9.6 8.5 8.8 8.7 8.9 8.0 Maxillary length 10.2 8.9 9.5 9.2 9.3 8.3 10.2 9.4 Eye diameter 7.5 7.1 7.3 7.8 7.6 8.5 8.7 8.0 Bony interorbital width 10.7 10.0 10.3 10.1 10.3 10.5 10.1 10.6 Depth of caudal peduncle 11.9 10.7 11.1 10.3 11.2 10.7 10.1 10.4 Length of caudal peduncle 17.7 18.3 15.8 17.9 17.8 19.2 16.9 18.8 Predorsal distance 50.3 50.9 50.4 50.8 50.9 50.2 50.9 49.9 Preanal distance 49.1 49.2 50.4 50.6 47.7 49.3 52.9 49.9 Prepelvic distance 37.8 36.6 39.7 38.3 35.9 37.1 40.0 40.2 2nd dorsal-fin base 25.8 21.7 23.8 22.6 24.7 23.8 21.8 20.2 Anal fin base 41.3 38.5 40.2 36.3 41.7 38.9 35.1 34.7 Pectoral-fin length 18.1 18.2 18.5 18.6 17.7 18.6 18.8 19.5 Pelvic fin-length 15.6 15.9 15.2 15.2 16.2 14.6 13.9 14.7 Longest ray 1st dorsal fin 16.2 12.0 11.1 11.3 14.5 14.8 11.6 11.6 Longest ray 2nd dorsal fin 13.1 12.3 11.1 10.9 11.8 10.7 11.4 12.3 Longest anal ray 18.2 13.2 13.3 13.0 14.9 13.1 12.7 13.7 Caudal-fin length 21.6 19.9 19.9 16.8 22.7 22.3 19.5 23.4 Caudal concavity 7.4 8.5 4.6 5.3 6.9 8.5 5.9 7.7 small scales basally on caudal fin and triangular longest (middle in both sexes) anal rays 1.5 (1.8- scale patch medially between base of pelvic fins; 2.2) in HL; pelvic-fin tips when depressed reaching predorsal scales extending forward to posterior half to base of second or third soft anal-fin ray in of interorbital space; preopercle with three scale mature males and females, but falling well short of rows between its posterior angle and eye. this point in immature fish; length of pelvic fins Predorsal length 2.0 (1.9-2.0) in SL; preanal 1.7 (1.6-2.0) in HL; length of pectoral fins 1.5 length 2.0 (1.9-2.1) in SL; prepelvic length 2.6 (1.3-1.5) in HL; length of caudal fin 1.3 (1.1-1.5) (2.5-2.8) in SL; length of second dorsal-fin base in HL; caudal-fin moderately forked, caudal con- 3.9 (4.0-4.9) in SL; length of anal-fin base 2.4 cavity 3.7 (3.0-5.8) in HL. (2.4-2.9) in SL. Colour of male holotype in life (Fig. First dorsal-fin origin about level with to slightly 2): blue grey with broad red scale margins, impart- behind anal-fin origin; longest spine (usually third) ing overall rich red appearance over most of body; of first dorsal fin 1.7 (1.8-2.5) in HL, its depressed head mainly grey blue except light greenish dor- tip reaching spine of second dorsal fin in females sally, grading to whitish ventrally with reddish and reaching to about base of second soft ray in chin; dorsal anal, and pelvic-fins dark brown to mature males; longest rays (middle in both sexes) blackish; second dorsal-fin with orange red outer of second dorsal fin 2.1 (2.1-2.6) in HL, depressed margin; caudal-fin pinkish orange; pectoral-fins posterior rays extending about one half of caudal translucent. peduncle in females and two-thirds to three- Colour of holotype in alcohol (Fig. 3): fourths length of caudal peduncle in mature males; after one month preservation largely orange red

141 aqua vol. 19 no. 3 - 19 July 2013 Melanotaenia sneideri, a New Species of Rainbowfish (Melanotaeniidae), from West Papua Province, Indonesia except uppermost portion of body brown; head (Fig. 4): brown on upper half of body, grading to dark grey except cheek and chin whitish; relatively whitish ventrally; broad (about two scale rows faint grey mid-lateral stripe on posterior third of wide), mid-lateral black stripe on posterior half of body, most conspicuous on caudal peduncle; dor- body; dorsal, anal, and pelvic-fins blackish, except sal-fins reddish brown, second dorsal fin with red- narrow white outer margin on second dorsal fin; dish outer margin; anal and pelvic-fins black; cau- opercle silvery grey with broad blackish area on dal-fin pale orange; pectoral-fins translucent uppermost portion; cheek brown; caudal and pec- whitish. The 69.6 mm SL male paratype is gener- toral-fins translucent whitish, caudal with dusky ally similar except the reddish hue has faded and grey rays on basal half. Small juveniles 15.1-27.4 the overall colouration is similar to that described mm SL, are either entirely whitish or pale grey on for the female below with the exception of the mid- the dorsal half of the body and white on the lower lateral blackish stripe, which extends forward to the half with a dark mid-lateral stripe that is vivid on upper margin of the opercle, although it is relatively the posterior one-third of the body (including cau- inconspicuous on the anterior half of the body. dal peduncle), but faint and inconspicuous on the Colour of adult female in alcohol anterior two-thirds.

Fig. 3. Melanotaenia sneideri, preserved male, holotype, 80,1 mm SL. Photo by G. R. Allen.

Fig. 4. Melanotaenia sneideri, preserved female, paratype (WAM P.33872-001), 79.4 mm SL. Photo by G. R. Allen. aqua vol. 19 no. 3 - 19 July 2013 142 Gerald R. Allen and Renny K. Hadiaty

Table II. Summary of dorsal, anal, and pectoral-fin ray, gill-raker, cheek-scale, and circumpeduncular-scale (CPS) counts for Melanotaenia sneideri. Pectoral-ray counts were taken on both sides of each individual.

First Dorsal Fin Spines Soft Dorsal Rays Soft Anal Rays IVVVI 1314 2324252628 2131 142 3 8 2 21

Pectoral Rays Total Gill Rakers CPS Cheek Scales 13 14 15 18 19 20 15 16 14 15 16 17 18 2237 8 6 1 5 8 4 2 52 1

Sexual dimorphism: Adult males are easily distin- with the longest rays in the middle portion of the guished on the basis of the overall red colouration, fin. This feature is shared by several members of which is lacking in females. Similar to most Melan- the genus from the Bird’s Head Peninsula includ- otaenia, males are also deeper bodied than females. ing M. ammeri Allen et al., 2008, M. irianjaya Unlike most members of the genus, which have a Allen, 1985, M. kokasensis Allen et al., 2008, and more elongate, pointed shape posteriorly on the M. parva Allen, 1990. Allen et al. (2008) and soft dorsal and anal fins of males, the male of M. Unmack et al. (2013) presented genetic evidence sneideri has a more evenly rounded anal-fin profile, indicating these allopatric species are closely related, forming a distinct subclade referred to as the “Southern Bird’s Head” group. Distribution and habitat: The type locality and only know location is situated in the Kumawa Mountains at the southern extremity of the Bomberai Peninsula of West Papua Province, Indonesia (Fig. 1). The location, which lies at an altitude of 1,050 m, consists of a small ephemeral lake basin, approximately 1000 m in length and 600 m wide. During dry periods it consists of small creek that emerges from limestone rocks (Fig. 5) on the edge of the basin and flows for 500-600 m before draining underground. The maximum width of this remarkably clear creek varies from about 1-2 m, with a maximum depth of 30-50 cm and average depth of about 10-20 cm. During a preliminary visit in late early March 2013, most fish, particularly large adults were concentrated in rocky pools near the end of the creek, just before it disappeared into the ground. However, conditions had changed dramatically due to heavy rainfall during a second visit near the end of March. The creek was then inundated, forming a small lake (Fig. 6) and the fish was consequently more widely dispersed. Etymology: The new species is named in honor of Richard Sneider, whom, together with Max Ammer planned and executed the 2013 Kumawa Mountains Expedition. Doctor Sneider discovered Fig. 5. Habitat of Melanotaenia sneideri at type locality, the species, photographed and filmed it, and, with showing origin of creek. Photo by R. Sneider. Obed Holago, first collected the type specimens.

143 aqua vol. 19 no. 3 - 19 July 2013 Melanotaenia sneideri, a New Species of Rainbowfish (Melanotaeniidae), from West Papua Province, Indonesia

Fig. 6. Habitat of Melanotaenia sneideri at type locality, showing formation of small lake after heavy rains. Photo by R. Sneider.

Fig. 7. Aquarium photograph of Melanotaenia parva, males, approximately 50 mm SL, Lake Kurumoi, West Papua Province, Indonesia. Photo by N. Khardina. aqua vol. 19 no. 3 - 19 July 2013 144 Gerald R. Allen and Renny K. Hadiaty

Remarks: Melanotaenia sneideri is separable from MZB 16455, 82.2 mm SL (holotype); AMS all other members of the genus and most melano- I.44640-001, 4 specimens, 46.5-66.5 mm SL taeniids on the basis of the unique male coloura- (paratypes); MZB 16456, 4 specimens, 55.9-71.5 tion. Only Glossolepis incisus Weber, 1907 from mm SL (paratypes); USNM 391630, 5 specimens, Lake Sentani, Papua Province (02°36.867’S, 46.1-71.6 mm SL (paratypes); WAM P. 33011- 140°33.743’E), possesses a substantial amount of 001, 5 specimens: 53.8-73.5 mm SL (paratypes); red on the body of adult males. Melanotaenia parva Melanotaenia irianjaya – MZB 4952, 50.0 mm SL from Lake Kurumoi, West Papua (2°09.528’S, (holotype), Fruata, 2°58.980’S, 133°31.977’E, 134°05.217’E) is also known to develop a bright Bomberai Peninsula; WAM P. 27863-001, 12 spec- red colour in captivity (Fig. 7), in contrast to the imens (paratypes), 20.0-51.0 mm SL, Fruata; mainly bluish to mauve hue of wild-caught fish. WAM P.29955-001, 10 specimens, 26.8-64.6 mm The adult male of M. sneideri is also separable from SL, Kali Satu, 2°05.913’S, 133°30.930’E, Bintuni all congeners by the dark brown to blackish colour vicinity; WAM P.29960-001, 53 specimens, 29.9- of the dorsal, anal, and pelvic fins. The maximum 78.1 mm SL, Kali Tujuh, 2°05.959’S, body depth (at least 42.3% of SL) attained by adult 133°29.988’E, Bintuni vicinity; Melanotaenia males further distinguishes it from most other kokasensis (all from vicinity of Kokas, 2°44.185’S, Melanotaenia from the Bird’s Head region, which 132°25.697’E) – MZB 16453, 56.5 mm SL (holo- generally have values less than 40%. Notable type); MZB 16454, 10 specimens, 27.2-52.9 mm exceptions are M. ajamarurensis Allen & Cross, SL (paratypes); USNM 391629, 7 specimens, 1980, M. angfa Allen, 1990, M. boesemani Allen & 36.2-64.0 mm SL (paratypes); WAM P. 33010- Cross, 1980, M. kamaka Allen & Renyaan, 1996b, 001, 7 specimens: 40.7-68.5 mm SL (paratypes); and M. lakamora Allen & Renyaan, 1996b. All of Melanotaenia parva – WAM P.29970-001, 125 these species except M. angfa inhabit lacustrine specimens (paratypes), 21.2-52.6 mm SL, Lake environments. Although there are a few exceptions, Kurumoi, 2°09.528’S, 134°05.217’E. lacustrine species of Chilatherina, Glossolepis, and Melanotaenia, generally have deeper bodied males ACKNOWLEDGEMENTS than stream-dwelling congeners. The more slender We are especially grateful to Max Ammer, owner shape of fish from lotic habitats is probably a of Sorido Bay Resort, Raja Ampat Islands, West hydro-dynamic adaptation to swift-flowing water. Papua, for informing us of this discovery and for- It is interesting to note that a deeper-bodied shape warding the specimens to the first author. We also is frequently assumed in aquarium-bred specimens thank Obed Holago of Wamena, Papua Province of lotic species compared to individuals of the same and Dr. Richard Sneider of Los Angeles, California species in nature (personal observations). for collecting the type series and providing pho- Although the largest collected specimen of M. tographs of the new species. Final thanks are due sneideri is 80.1 mm SL, the species attains a much Tertius Kammeyer, who assisted with expedition larger size. One specimen, 110 mm SL was col- logistics and provided locality information. lected, but unfortunately discarded due to the lack of a sufficient-sized preservation container. This is one of the three largest known species from the REFERENCES Bird’s Head Peninsula. Only M. fasiensis Kadarus- ALLEN, G. R. 1985. Three new rainbowfishes (Melano- man et al., 2010, (120 mm SL) and M. angfa (114 taeniidae) from Irian Jaya and Papua New Guinea. Revue française d’Aquariologie Herpétologie 12 (2): 53-62. mm SL) are known to attain a larger size. ALLEN, G. R. 1990. Les poissons arc-en-ciel (Melanotaeni- Other noteworthy features that further distin- idae) de la Péninsule de Vogelkop, Irian Jaya, avec guish the new species from most Melanotaenia description de trois nouvelles espèces. Revue française include the combination of 15-16 circumpeduncu- d’Aquariologie Herpétologie 16 (4) (1989): 101-112. lar scales and 18-20 total gill rakers on the first ALLEN, G. R. 1995. Rainbowfishes in Nature and in the branchial arch, both values that are relatively high Aquarium. Tetra Verlag, Melle, Germany, 180 pp. for the genus compared to counts of 11-14 and 13- ALLEN, G. R. 1996. Two new species of rainbowfishes 18 respectively for most other species. (Melanotaenia: Melanotaeniidae), from the Kikori River system, Papua New Guinea. Revue française d’Aquariolo- Comparative material (all from West Papua gie Herpétologie 23 (1-2): 9-16. Province, Indonesia): Melanotaenia ammeri (all ALLEN, G. R. 1997. A new species of rainbowfish (Melano - from Arguni Bay, 3°02.438’S, 133°52.844’E) – taenia: Melanotaeniidae), from the Lakekamu Basin,

145 aqua vol. 19 no. 3 - 19 July 2013 Melanotaenia sneideri, a New Species of Rainbowfish (Melanotaeniidae), from West Papua Province, Indonesia

Papua New Guinea. Revue française d’Aquariologie Herpé- U.S. Geological Survey Professional Paper 1078. U.S. tologie 24 (1-2): 37-42. Government Printing Office, Washington, D.C. ALLEN, G. R. 1998. A new genus and species of rainbow- HERRE, A. W. C. T. 1935. New fishes obtained by the fish (Melanotaeniidae) from fresh waters of Irian Jaya, Crane Pacific expedition. Field Museum of Natural His- Indonesia. Revue française d’Aquariologie Herpétologie 25 tory, Publications, Zoölogical Series 18 (12): 383-438. (1-2): 11-16. HILL, K. C. & HALL, R. 2003. Mesozoic-Cenozoic evolu- ALLEN, G. R. 2001. A new species of rainbowfish (Glos- tion of Australia’s New Guinea margin in a west Pacific solepis: Melanotaeniidae) from Irian Jaya, Indonesia. Jour- context. In: Evolution and Dynamics of the Australian nal of the Australia New Guinea Fishes Association 15 (3): Plate (Eds. R.R. Hillis, & R.D. Müller): 265-290. Geo- 766-775. logical Society of Australia Special Publication 22 and ALLEN, G. R. & CROSS, N. J. 1980. Descriptions of five Geological Society of America Special Paper 372. new rainbowfishes (Melanotaeniidae) from New Guinea. KADARUSMAN, HADIATY, R. K., SEGURA, G., SETIAWIBAWA, Records of the Western Australian Museum 8 (3): 377-396. G., CARUSO, D. & POUYAUD, L. 2012. Four new species ALLEN, G. R. & HADIATY, R. K. 2011. A new species of of rainbowfishes (Melanotaeniidae) from Arguni Bay, rainbowfish (Melanotaeniidae), from western New West Papua, Indonesia. Cybium 36 (2): 369-382. Guinea (West Papua Province, Indonesia). Fishes of Sahul KADARUSMAN, SUDARTO, PARADIS, E. & POUYAUD, L. 25 (1): 602-607. 2010. Description of Melanotaenia fasinensis, a new ALLEN, G. R., MIDGLEY, S. H. & ALLEN, M. 2002. Field species of rainbowfish (Melanotaeniidae) from West Guide to the Freshwater Fishes of Australia. Western Aus- Papua, Indonesia with comments on the rediscovery of tralian Museum, Perth, 394 pp. M. ajamaruensis and the endangered status of M. parva. ALLEN, G. R. & RENYAAN, S. J. 1996a. Chilatherina pricei, Cybium 34 (2): 207-215. a new species of rainbowfish (Melanotaeniidae) from KADARUSMAN, SUDARTO, SLEMBROUCK, J. & POUYAUD, L. Irian Jaya. Revue française d’Aquariologie Herpétologie 23 2011. Description of Melanotaenia salawati, a new (1-2): 5-8. species of rainbowfish (Melanotaeniidae) from Salawati ALLEN, G. R. & RENYAAN, S. J. 1996b. Three new species Island, West Papua, Indonesia. Cybium 35 (3): 223-230. of rainbowfishes (Melanotaeniidae) from the Triton MCGUIGAN, K. L. 2001. An addition to the rainbowfish Lakes, Irian Jaya, New Guinea. aqua, Journal of Ichthyo- (Melanotaeniidae) fauna of north Queensland. Memoirs logy and Aquatic Biology 2 (2): 13-24. of the Queensland Museum 46 (2): 647-655. ALLEN, G. R. & RENYAAN, S. J. 1998. Three new species PIGRAM, C. J. & DAVIES, H. L. 1987. Terranes and the of rainbowfishes (Melanotaeniidae) from Irian Jaya, accretion history of the New Guinea orogen. Bureau of Indonesia. aqua, Journal of Ichthyology and Aquatic Bio- Mineral Resources, Journal of Australian Geology and Geo- logy 3 (2): 69-80. physics 10: 193-211. ALLEN, G. R. & UNMACK, P. J. 2008. A new species of POLHEMUS, D. A. 2007. Tectonic geology of Papua. In: rainbowfish (Melanotaeniidae: Melanotaenia) from The Ecology of Papua. (Eds. A. J. Marshall & B. M. Batanta Island, western New Guinea. aqua, International Beehler): 137-164. Periplus Editions, Singapore. Journal of Ichthyology 13 (3-4): 109-120. PRICE, D. S. 1997. Chilatherina alleni, a new species of ALLEN, G. R. & UNMACK, P. J. 2012. A new species of rainbowfish (Melanotaeniidae) from Irian Jaya. Revue rainbowfish (Chilatherina: Melanotaeniidae) from the française d’Aquariologie Herpétologie 24 (3-4): 79-82. Sepik River system of Papua New Guinea. aqua, Interna- RAMSAY, E. P. & OGILBY, J. D. 1886. A contribution to tional Journal of Ichthyology 18 (4): 227-237. the knowledge of the fish-fauna of New Guinea. Proceed- ALLEN, G. R., UNMACK, P. J. & HADIATY, R. K. 2008. Two ings of the Linnean Society of New South Wales (Series 2) 1 new species of rainbowfishes (Melanotaenia: Melano- (1): 8-20. taeniidae), from western New Guinea (Papua, Barat UNMACK, P. J., ALLEN, G. R. & JOHNSON, J. B. 2013. Phy- Province, Indonesia). aqua, International Journal of logeny and biogeography of rainbowfishes (Melanotaeni- Ichthyology 14 (4): 209-224. idae) from Australia and New Guinea. Molecular Phyloge- CASTELNAU, F. L. 1875. Researches on the fishes of Aus- netics and Evolution 67: 15-27. tralia. Philadelphia Centennial Expedition of 1876. Inter- WEBER, M. 1907. Süsswasserfische von Neu-Guinea. Ein colonial Exhibition Essays, 1875-6: 1-52. Beitrag zur Frage nach dem früheren Zusammenhang GRAF, J. & OHEE, H. L. 2009. Glossolepis from northern von Neu-Guinea und Australien. In: Nova Guinea. Résul- New Guinea. Fishes of Sahul. Journal of the Australian tats de l’expédition scientifique Néerlandaise à la Nouvelle- New Guinea Fishes Association 23 (2): 502-512. Guinée en 1903. (Ed. A. Wichmann): Volume 5 (Zool- HAMILTON, W. B. 1979. Tectonics of the Indonesian region. ogy) part 2: 201-267. E. J. Brill, Leiden.

aqua vol. 19 no. 3 - 19 July 2013 146 aqua, International Journal of Ichthyology

A new species of Egglestonichthys from northern Australian estuaries (Teleostei, Gobiidae, Gobiinae)

Helen K. Larson

Museum and Art Gallery of the Northern Territory, P.O. Box 4646, Darwin, NT 0801, Australia; Museum of Tropical Queensland, 70-102 Flinders street, Townsville, Queensland 4810, Australia. School of Marine and Tropical Biology, James Cook University, Townsville, Queensland 4811, Australia.

Received: 02 June 2013 – Accepted: 07 July 2013

Abstract midae). L’espèce la plus proche pourrait être E. bombylios A new species, that appears to belong to the gobiid genus Larson et Hoese, bien que le patron de ses papilles senso- Egglestonichthys, is described from specimens collected from rielles diffère de celles des autres membres du genre. Une the shallow coast and estuaries of Kakadu National Park in clé des quatre espèces d’Egglestonichthys connues à ce jour the Northern Territory, Australia. It is unusual among gobi- est fournie. ids in that the lower half of the pectoral-fin lacks connecting membrane between the fin rays, so that the rays are free Sommario (resembling the pectoral fin of the fishes of the family Una nuova specie, che sembra appartenere ai gobidi del Polynemidae). Its closest relative may be E. bombylios Lar- genere Egglestonichthys, è descritta sulla base di esemplari rac- son and Hoese, although its sensory papilla pattern differs colti in estuari e zone costiere poco profonde del Kakadu from others in the genus. A key to the four known species National Park nel Northern Territory, Australia. Presenta presently placed in Egglestonichthys is provided. un’insolita caratteristica tra i gobidi: la metà inferiore della pinna pettorale manca di membrana di collegamento tra i Zusammenfassung raggi, che, pertanto, risultano liberi. (ciò la rende simile alla Beschrieben wird eine neue Art, die offenbar zu den pettorale dei pesci della famiglia Polynemidae). Il suo Grundeln der Gattung Egglestonichthys gehört, nach parente più prossimo potrebbe essere E. bombylios Larson e Exemplaren, die an der flachen Küste und in den Münd- Hoese, anche se il modello di distribuzione delle papille ungsgebieten im Kakadu-Nationalpark des Nordterritori- sensoriali è diverso da tutte le altre specie del genere. Viene ums in Australien gesammelt wurden. Sie zeigen ein unter fornita una chiave di identificazione per le quattro specie Gobiiden ungewöhnliches Merkmal: die untere Hälfte der attualmente conosciute collocate in Egglestonichthys. Brustflossen hat keine verbindende Membran zwischen den Flossenstrahlen, sodass die Strahlen frei stehen (ähnlich wie INTRODUCTION bei den Fischen der Familie Polynemidae). Als nächster In 1997, during an estuarine fish survey along the Verwandter dürfte E. bombylios Larson and Hoese gelten, obwohl das Muster der Sinnes-Papillen sich von allen ande- coast of Kakadu National Park, Northern Territory, ren in der Gattung unterscheidet. Beigefügt ist ein Bes- specimens of an unusual small gobiid were obtai- timmungsschlüssel zu den vier Arten, die bisher der Gatt- ned by a small beam trawl (2 m wide mouth, 6.5 ung Egglestonichthys zugeordnet werden konnten. mm mesh body and 1.4 mm mesh cod end). Subsequent surveys in 1998, 1999 and 2001 Résumé produced additional specimens. The fish have been Une nouvelle espèce, qui semble appartenir au genre de tentatively identified as belonging to Egglestonich - Gobiidés Egglestonichthys, est décrite sur base de spécimens thys, a genus containing three other species of collectés sur la côte étroite et dans les estuaires du parc somewhat disparate appearance. Miller and Won- national de Kakadu, Territoire du Nord, Australie. Elle diffère des autres Gobiidés par le fait que la moitié infé- grat (1979) created the genus for a goby with a rieure des pectorales n’a pas de membranes entre les complex transverse papilla pattern resembling that rayons, ce qui implique que ces nageoires sont libres (évo- found in some eleotrids (e.g. Eleotris), lacking late- quant la pectorale des poissons de la famille des Polyne- ral-line sensory pores and having a wide gill open-

147 aqua vol. 19 no. 3 - 19 July 2013 A new species of Egglestonichthys from northern Australian estuaries (Teleostei, Gobiidae, Gobiinae) ing (to below preoperculum). Larson and Hoese The new species somewhat resembles E. bombylios (1997) described a second species (E. bombylios) Larson and Hoese, 1997, in general appearance, and redescribed E. melanoptera Rao, originally having reduced eyes, moderate gill opening and described as a Callogobius. Miller and Wongrat with some raised sensory papilla rows on the head, (1979) provided the only published osteological but differing in having a number of fin rays in the information for the genus (based on the type spe- lower half of the pectoral-fin free from connecting cies, E. patriciae). membranes, resembling the rays of a polynemid,

Fig. 1. Locality map for known records of Egglestonichthys ulbubunitj n. sp. Map prepared by Michael Hammer. aqua vol. 19 no. 3 - 19 July 2013 148 Helen K. Larson no scales on the cheek or opercle, as well as a sen- Museum, Sydney; NTM, Museum and Art Gallery sory papilla arrangement that is unlike other spe- of the Northern Territory, Darwin: QM, Queens- cies of Egglestonichthys. Although the new species land Museum, Brisbane; WAM, Western Aust- has several features that differ from other Egglesto- ralian Museum, Perth. nichthys, the group is so little-studied that I am reluctant to create yet another new genus based on limited information. Egglestonichthys ulbubunitj n. sp. So far this fish is known only from coastal marine Threadfin Goby and estuarine waters within Kakadu National Park (Figs 2-4, Table I) (Larson 2002) (Fig. 1). Similar beam trawl and dredging surveys have been carried out in other Holotype: NTM S.14635-005, 28 mm SL female, estuaries and harbours in the Northern Territory, off Pococks Beach, N of West Alligator Head, 1.4 but this species has not been not found. It is most m, very soft mud, beam trawl, 26 May 1998, R. interesting that this new species has not yet been Williams, G. Dally, A. Pickworth and M. Gorst. collected elsewhere in the Territory, and it is pos- Paratypes: NTM S.17348-001, 3(26.5-30), same sible that the fish has very specific habitat require- data as holotype; NTM S.14479-003, 1(31), ments. centre of West Alligator River mouth, 5.5 m, mud, beam trawl, 27 May 1997, coll. R. Williams and A. METHODS Pickworth; NTM S.14657-007, 4(10.5-20), off Measurements were taken using electronic calli- “The Rookery”, East Alligator River, 7.3 m, mud, pers and dissecting microscope. Counts and met- beam trawl, 2 June 1998, coll. R. Williams and G. hods generally follow Hubbs and Lagler (1970), Lindner; NTM S.15703-001, 1(18.5), 5 km offs- except as indicated below. Transverse scale counts hore between Field Island and Point Farewell, 9.6 backward (TRB) are taken by counting the num- m, mud, beam trawl, coll. R. Williams, S. Morr- ber of scale rows from the anal-fin origin diago- ison and D. Elphick, 12 June 2001; NTM nally upward and back toward the second dorsal- S.16024-001, 1(28), N of West Alligator Head, 16 fin base. Head length is taken to the upper attach- m, soft mud and shell grit, dredge, coll. N. Smit ment of the opercular membrane. References to and party, 21 November 2004; QM I.39101, second dorsal and anal-fin rays includes a thin flex- 4(18-24), 1 km off Pococks Beach, West Alligator ible spine in addition to the segmented rays. The Head, 5 m, mud, beam trawl, coll. A. Pickworth, segmented or branched caudal ray pattern (e.g. 9/8 M. Gorst and A. Turner, 7 June 1999; AMS or 9/7) is the number of segmented caudal rays I.46190-001, 2(24-27), off Midnight Point, South attaching to the upper and lower hypural plates Alligator River, 10 m, mud and detritus, beam respectively. Vertebral counts and other osteologi- trawl, coll. R. Williams, S. Morrison and D. Elp- cal information were obtained by X-ray. Pterygiop- hick, 12 June 2001; NTM S.15289-003, 1(19), 31 hore formula follows Birdsong et al. (1988). km upstream from mouth of South Alligator River, Institutional abbreviations are: AMS, Australian 10-11 m, mud, beam trawl, coll. G. Lindner, F.

Fig. 2. Holotype of Egglestonichthys ulbubunitj n. sp., NTM S.14635-005, 28 mm SL female. Photo by J. Larson.

149 aqua vol. 19 no. 3 - 19 July 2013 A new species of Egglestonichthys from northern Australian estuaries (Teleostei, Gobiidae, Gobiinae)

Table I. Morphometrics of Egglestonichthys ulbubunitj n. sp., expressed as percentage of SL or HL as indicated (n = 21).

Holotype Mean Maximum Minimum

Head length in SL 29.3 30.4 33.3 27.9 Head depth in HL 69.5 67.7 75.0 61.7 Head width in HL 76.8 68.5 83.0 60.7 Body depth in SL 23.9 21.2 23.9 18.4 Body width in SL 9.3 8.6 10.4 7.0 Caudal peduncle length in SL 19.6 20.5 24.6 18.3 Caudal peduncle depth in SL 13.6 11.8 13.6 10.5 Snout length in HL 23.2 23.5 27.7 20.0 Eye width in HL 15.9 15.9 18.3 13.5 Upper jaw length in HL 42.7 45.5 50.0 38.3 Interorbital width in HL 23.2 18.4 23.2 14.0 Pectoral-fin length in SL 27.9 29.0 33.7 26.7 Pelvic-fin length in SL 28.6 29.5 33.9 26.7 Caudal-fin length in SL 53.6 56.2 63.6 50.0 Longest dorsal-fin spine in SL 13.6 15.9 19.1 13.0

Baird and G. McSkimming, 6 June 1999; WAM P. South Alligator River, 12° 12’ 48.5994” 132° 24’ 33873-001, 1(33), Brooks Creek mouth, South 53.9994”, in 3.5m depth over mud, beam trawl, Alligator River, 8-9 m, mud, beam trawl, coll. G. coll. B. Pusey and M. Kennard, 27 August 2012 Lindner, F. Baird and G. McSkimming, 6 June [frozen in the field for later identification]. 1999; NTM S.14441-012, 2(20-24.5), off mouth Diagnosis: A small, slender, round-headed goby of Wildman River, 2 m, firm mud and plant detri- with small eyes, long pointed caudal fin and lower tus, beam trawl, coll. H. Larson, R. Williams, A. 3-7 pectoral-fin rays free from membrane; second Pickworth and M. Gorst, 28 May 1997. dorsal-fin rays always I,10; anal fin rays always Non-type material: NTM S.17479-001, 1(29), I,10; pectoral-fin rays 19-21; longitudinal scales

Fig. 3. Egglestonichthys ulbubunitj n. sp., diagram of head showing sensory papillae, NTM S.15703-001, 18.5 mm SL female. aqua vol. 19 no. 3 - 19 July 2013 150 Helen K. Larson

22-25; TRB 7-9; side of head naked, predorsal sca- 15*), 3-2310 (in 1). Vertebrae 10+16 (in 18). One les 9-12, extending forward to behind eyes; eyes epural (in 17*), two in one specimen. Two anal small and dorsolateral; interorbital broad and con- pterygiophores before haemal spine of first caudal vex; jaws oblique, with small caniniform teeth; vertebra (in 18). head pores absent; reduced papillae pattern on Head rounded in cross-section, length 27.9- head arranged in mostly longitudinal pattern, 33.3% (mode 30.0%) in SL; profile rounded to papillae often pointed, fleshy and may be on short slightly pointed. Depth at posterior preopercular raised ridges; eyes silver when alive, head and body margin 61.7-75.0% (mode 66.7%) in HL. Head colour pale yellowish-white, with six or seven ind- width at posterior preopercular margin 60.7- istinct dusky bands across nape and sides, and large 83.0% (mode 76.8%) in HL. Mouth terminal, oval black spot on posterior part of first dorsal fin. oblique, forming an angle of 35-40° with body Description: Based on 21 specimens, 18-33 mm axis; jaws reaching to below posterior margin of eye SL. Counts of holotype (Fig. 2), where differing or at least posterior half. Lips smooth, lower lip from paratypes, indicated by asterisk (in parenthe- free, fused on either side of broad flat chin. Upper ses where necessary); morphometrics in Table I. jaw 38.3-50.0% (mode 45.0%) in HL. Eye very First dorsal fin VI; second dorsal-fin rays always small, placed laterally, closer to snout tip than to I,10; anal-fin rays always I,10; pectoral-fin rays 19- rear margin of preopercle, 13.5-18.3% (mode 21 (usually 20 (holotype with 20 in left fin, 21 on 15.9%) in HL. Snout broad and short, 20.0- right); segmented caudal-fin rays 17; caudal-fin ray 27.7% (mode 23.2%) in HL. Interorbital very pattern 9/8 (one specimen with 9th upper ray split broad, flattened, 14.0-23.2%% (mode 19.0%) in to near base); branched caudal-fin rays 8/6 (in 10), HL. Body slender and compressed, depth at anal 8/7* (in 11), 7/6 (in 1); unsegmented (procurrent) origin 18.4-23.9%% (mode 23.9%) in SL; body caudal-fin rays 6/6 to 8/8 (usually 7/6 or 7/7, 8/7 width at origin of first dorsal 7.0-10.4% (mode in holotype); longitudinal scale count 22-25 7.0%) in SL. Caudal peduncle length 20.0-24.6% (modally 23*); TRB 7-9 (modally 8*); predorsal (mode 20.0%) in SL. Caudal peduncle depth 10.5- scale count 9-12 (modally 9; 11 in holotype); circ- 13.6% (mode 13.6%) in SL. umpeduncular scales 12*-13 (modally 12). Gill No mental fraenum; single transverse row of rakers on outer face of first arch 3+11 (1), 4+11 fleshy papillae present on chin. Anterior naris in (1), 5+11 (1). Pterygiophore formula 3-22110* (in broad, very short tube, close behind upper lip.

Fig. 4. Egglestonichthys ulbubunitj n. sp., showing fresh dead colour pattern; photograph taken through microscope; whole fish could not be viewed; NTM S.17479-001, 29 mm SL, South Alligator River. N.B. some small yellowish patches of org- anic material still adhere to the specimen. Photo courtesy of Charles Darwin University.

151 aqua vol. 19 no. 3 - 19 July 2013 A new species of Egglestonichthys from northern Australian estuaries (Teleostei, Gobiidae, Gobiinae)

Posterior naris rounded to oval, placed above and ral-fin rays branched, upper and lowermost rays forward of anterior margin of eye. Gill opening unbranched; lowermost three to seven (usually five moderate, extending forward to just under opercle to six, three in only one specimen) rays free from or to mid-opercle, but not reaching posterior preo- membrane (tips of branches of each individual percular margin. Anterior edge of shoulder girdle branched ray joined by membrane). Pelvic-fins (cleithrum) smooth. Gill rakers on outer face of large (reaching anus), oval, disc-shaped, fraenum first arch without spines, fleshy and slender, deep; fin length 26.7-33.9% (mode 28.6%) in SL. somewhat shorter than gill filaments; rakers on Caudal fin very long and pointed, 50.0-63.6% inner face of first arch short, stubby, without fine (mode 53.6%) in SL. spines; outer rakers on other arches similar to inner Live coloration: Only a few notes were face rakers. Tongue short, blunt to slightly taken when the holotype was collected: “translucent rounded. grey fat-headed goby with small silver eyes and ind- Outer row teeth in upper jaw small, sharp and istinct grey cross-hatched bands across back”. Fig- pointed, in three rows across front and two along ure 4 shows the freshly dead non-type specimen. side; outermost row teeth larger, more curved, with Coloration in alcohol: Background largest two to six teeth on either side of symphysis. colour yellowish white, with six or seven short faint Outer row teeth in female slightly smaller than brownish saddles crossing back; first on nape those in male. Lower jaw with two to three rows of similarly sized, small sharp teeth, none particularly large; outermost row teeth may be more upright and less curved. Scales on head and body all cycloid. Predorsal scales small, reaching forward to close behind eyes. Cheek and operculum without scales. Pectoral base naked or with few large scales. Prepelvic area with moderately-sized scales, may be embedded. Head pores absent. Sensory papillae pattern on head in pattern as shown diagrammatically in Figure 3; appearing longitudinal but with few rows that appear transverse (and with some individual variation); all specimens show some damage to sensory papillae due to abrasion during trawling and subsequent handling. Papillae on head large, fleshy and often with pointed tips, in some specimens, papillae appear as two forms: vertically oriented papillae on short raised ridges, horizontally oriented papillae small and oval to round, within rounded fleshy depressions. In some specimens, papillae barely visible above skin, and can be seen through partly transparent flesh as short ridges; papillae on under- side of head usually discernible above skin. Genital papilla in male short, small, flattened, with blunt to slightly pointed tip. Genital papilla in females very short, bulbous. First dorsal-fin low, rounded, spines barely reaching first element of second dorsal-fin origin; third or fourth dorsal spine longest, 13.0-19.1% in SL. Fig. 5. Typical beam-trawl catch on the South Alligator Second dorsal and anal-fins long, pointed posteri- River, Northern Territory, after 20 minutes’ haul with 2 orly, posteriormost rays of these fins usually reach- metre beam-trawl. Rex Williams is holding an adult Strea- ing onto caudal fin. Pectoral-fin relatively narrow, mer Threadfin, Parapolynemus verekeri (Polynemidae), pointed, 26.7-33.7% (mode 30.0%) in SL; pecto- which co-occurs with E. ulbubunitj. Photo by H. K. Larson. aqua vol. 19 no. 3 - 19 July 2013 152 Helen K. Larson before first dorsal fin and seventh at caudal-fin base nal Owner Jonathan Nadji. He stated: “My family’s (two saddles under second dorsal fin may coalesce); clan area includes the area where this fish (goby saddles usually joining short staggered oblique pale threadfin) was discovered. Our clan, Ulbu Bunidj, brownish bars or blotches along midside of body; shares this area with Djindibi. I would like this fish within brownish saddles, scale margins narrowly to be named after our clan, which is Ulbu Bunitj”. outlined with slightly darker pigment (Fig. 4). Suggested common name: Threadfin Goby. Head and chin evenly speckled with fine brown; Remarks: This species will not key out to Egg- nape paler, may show one to three indistinct lestonichthys in Larson and Murdy’s 2001 key to brownish bars crossing from opercle to opercle. western Pacific gobiid genera. In fact, anyone Indistinct short brown line from front margin of attempting to use this key for any species of Egg- eye to middle to upper jaw. Remainder of under- lestonichthys will not succeed as couplet 35b is side of head, breast and belly whitish. incorrect, unfortunately (it states that head pores First dorsal-fin translucent, anteriorly with faint are present, when this is not so in Egglestonichthys; brownish wedge-shaped mark extending up from writing a working key to 76 often poorly defined second dorsal saddle; distinct black round to oval genera is difficult). spot on posterior part of fin extending from fourth As noted by Larson and Hoese (1997), there is to sixth spine. Second dorsal-fin translucent to some variation of features among the described faintly brownish, fin margin narrowly unpigmen- species of this genus. Egglestonichthys ulbubunitj ted. Anal-fin translucent, with variable amounts lacks spines on the gill rakers, as does E. bombylios. and intensity of brown speckling along basal half of The sensory papillae pattern on the head of E. fin, usually more prominent on posterior part of ulbubunitj is not like that of any of the other spe- fin. Pectoral-fin translucent, with faint wedge of cies, in that the transverse pattern is reduced and brown speckling on upper half, an extension of an varies somewhat between specimens, although the indistinct brown bar along bases of pectoral-fin papilla structures themselves (fleshy, partly raised rays and rear part of pectoral-fin base. Pelvic-fins on ridges) do resemble those of E. bombylios. Of translucent. Caudal-fin translucent with faint the four known species of the genus, only E. pat- brownish speckling basally and often along riciae and E. melanoptera are similar to each other membranes (mostly on lower half of fin). (and could be conspecific; fresh, undamaged Distribution: Known only from turbid estuarine material is needed). Neither E. bombylios nor E. and coastal habitats in the Alligator Rivers region, ulbubunitj greatly resemble the other two. The Northern Territory. paucity of specimens makes further analysis diffi- Comparisons: This is a distinctive fish, and is cult – one known specimen of E. patriciae, four unlikely to be confused with any other gobiid, each of E. melanoptera and E. bombylios as comp- especially when the pectoral fins are examined. It ared to 21 of E. ulbubunitj, the smallest species does not closely resemble the other species of the (N.B. have just learned that 30 specimens of E. genus Egglestonichthys; indeed, all four described melanoptera have recently been collected off India; species are rather different from each other in body pers. comm. D. Ray, Zoological Survey of India, form and colour pattern (see Remarks and dicho- West Bengal). It is possible that more specimens tomous key below). are waiting among the trawled fish identified only Ecology: All specimens were obtained by 2-metre as “Gobiidae” in museums. beam trawl or small dredge over (and occasionally Further material of all species will confirm whet- in) very soft to firm mud (occasionally with broken her these four odd goby taxa truly belong together shell and plant detritus such as drifts of leaves), at as a group or not. Genetic analyses and further depths of 1-16 m. Water quality varied with the morphological examination will undoubtedly help tide, from 25.8-28.4°C and 28.0-30.3‰ salinity. disentangle them. Although it is not entirely satis- This small goby co-occurs with estuarine fishes factory to assign this new species to a genus to such as small polynemids, juvenile sciaenids, which it may not belong, it at least flags the pro- engraulids and cynoglossids in this soft-substrate blem and indicates that much work remains to habitat (Larson 1999). be done on all those small “trawl gobies” presently Etymology: Named ulbubunitj, to be treated as a waiting to be examined in collections world-wide. noun in apposition, for the Ulbu Bunidj clan in Arnhem Land, at the suggestion of senior Traditio-

153 aqua vol. 19 no. 3 - 19 July 2013 A new species of Egglestonichthys from northern Australian estuaries (Teleostei, Gobiidae, Gobiinae)

Key to the species of Egglestonichthys with us in collecting these peculiar fish during the Kakadu National Park estuarine fish monitoring 1. Pelvic frenum present, may be thin (and easily surveys from 1997 to 2002. Their willingness and damaged); body pale to brownish with darker good humour made the difficult task of beam- bands or blotches; mouth small to large ……2 trawling in deep mud habitats that much more 1a. Pelvic frenum absent; fins and body generally enjoyable. plain dark brown to black; mouth large and reaching to well below eye...... E. melanoptera (Rao, 1971) REFERENCES 2. Lower 3-7 pectoral-fin rays free from fin BIRDSONG, R. S., MURDY, E. O. & PEZOLD, F. L. 1988. A membrane; caudal-fin elongate and pointed, study of the vertebral column and median fin osteology much longer than head; lateral scales 22-25 ... in gobioid fishes with comments on gobioid relations- ...... E. ulbubunitj n. sp. hips. Bulletin of Marine Science 42 (2): 174-214. 2a. All pectoral-fin rays contained within mem- HUBBS, C. L. & LAGLER, K. F. 1970. Fishes of the Great brane; caudal fin not elongate and pointed; Lakes Region. University of Michigan Press: Ann Arbor. with at least 30 lateral scales or more ...... 3 LARSON, H. K. 1999. Report to Parks Australia North, on the estuarine fish inventory of Kakadu National Park, 3. Papilla rows on head raised on fleshy ridges Northern Territory, Australia. Museum and Art Gallery especially around jaws, anterior cheek and of the Northern Territory Research Report No. 5. 50 pp. snout; lateral scales 31-35; eyes small (9-13 in LARSON, H. K. 2002. Report to Parks Australia North, on HL)..... E. bombylios Larson and Hoese, 1997 estuarine fish monitoring of Kakadu National Park, 3a. Papilla rows on head not raised on fleshy rid- Northern Territory, Australia. Museum and Art Gallery ges; lateral scales 40; eyes moderate (3.5 in of the Northern Territory, unpublished report. 42 pp. HL).... E. patriciae Miller and Wongrat, 1979 LARSON, H. K. & HOESE, D. F. 1997. A new species of Egglestonichthys (Pisces; Teleostei; Gobiidae) from the Indo-West Pacific, with discussion of the species of the ACKNOWLEDGEMENTS genus. The Beagle, Records of the Museums and Art Galler- My many thanks to Rex Williams and Gavin ies of the Northern Territory 13: 45-52. Dally of the Museum and Art Gallery of the Nort- LARSON, H. K. & MURDY, E. O. 2001. Gobiidae. Gobies. hern Territory for their considerable help. Thanks Pp 3578-3603. In: Carpenter, K. E. and Niem, V. H. to Charles Darwin University staff for photograp- (eds) FAO species identification guide for fishery purposes. hing the freshly dead colouring of the small non- The living marine resources of the western Central Pacific. paratype specimen. Many thanks also to Garry Volume 6. Bony fishes part 4 (Labridae to Latimeriidae). Linder of Parks Australia North for all his assist- FAO, Rome. MILLER, P. J. & WONGRAT, P. 1979. A new goby (Teleos- ance, especially with logistics; Parks Australia tei: Gobiidae) from the South China Sea and its signific- North staff, in particular Mick Gorst, Anna Pick- ance for gobioid classification. Zoological Journal of the worth, Buck Salau and Andrew Wellings, and Linnean Society 67: 239-257. Traditional Owners Joseph Bishop, Leanne Alder- RAO, V. V. 1971. New gobioids from Godavari estuary. son and Kenny Wauchope; all of whom worked Journal of the Zoological Society of India 23 (1): 39-54.

aqua vol. 19 no. 3 - 19 July 2013 154 aqua, International Journal of Ichthyology

New Distribution Records for the Cairns Rainbowfish Cairnsichthys rhombosomoides (Melanotaeniidae): implications for conservation of a restricted northern population

Keith C. Martin and Susan Barclay

P.O. Box 520 Clifton Beach Qld 4879 Australia. Email: [email protected]

Received: 10 May 2013 – Accepted: 08 July 2013

Abstract gestellten Bestände von C. rhombosomoides sind wegen ihrer Four new locality records for the Cairns Rainbowfish stark spezialisierten Habitatansprüche und der räumlich Cairnsichthys rhombosomoides Nichols & Raven, 1928 are begrenzten Verbreitung ausgesprochen empfindlich gegen documented. The new localities are all outside the known lokale Störungen, besonders solche, die den Fluss des geographical range for this species. Three of the new locali- Fließgewässers einschränken, wie vermehrte Trockenheit ties appear to represent part of a disjunct northern popula- und menschliche Maßnahmen, vor allem die Entnahme von tion in a biogeographical province where the species was Oberflächen- und Grundwasser zur Trinkwassergewinnung. previously unknown. The fourth locality appears to be a minor range extension of the existing known population. Résumé The new localities all contain habitats similar to those Quatre nouvelles localisations de Cairnsichthys rhomboso- found in areas where the core populations of C. rhombosomoides (Nichols & Raven 1928) sont documentées. Ces nou- moides occur and environmental data is provided for all of velles localisations sont toutes situées en dehors de la distri - the newly recorded localities. Specimens from the northern bution géographique connue de cette espèce. Trois des nou- localities are genetically distinct, although they appear mor- velles localisations semblent représenter une partie d’une phologically similar to those from the core area of distribu- population septentrionale séparée dans une province biogéo- tion. Cairnsichthys rhombosomoides populations at the new graphique où l’espèce était inconnue à ce jour. La quatrième localities have extremely restricted habitat requirements and localisation semble être une extension mineure de la popula- localised distributions, and therefore may be particularly tion existante connue. Les nouvelles localisations contien- vulnerable to local perturbations, especially those that may nent toutes des habitats semblables à ceux des régions où vi- reduce stream flows such as extended droughts and human vent les principales populations de C. rhombosomoides et des activities such as surface and ground water extraction. don nées environnementales sont fournies pour toutes les nouvelles localisations. Les spécimens des localisations sep - Zusammenfassung ten trionales sont génétiquement distincts, bien qu’ils sem- Dokumentiert werden vier neue Nachweise zum Vorkom- blent morphologiquement similaires à ceux de la distribution men des Regenbogenfisches Cairnsichthys rhombosomoides principale. Les populations de Cairnsichthys rhombosomoides (Nichols & Raven 1928). All diese vier neuen Nachweise des nouvelles régions ont des besoins en habitat extrê mement liegen außerhalb der bisher bekannten geographischen Ver- précis et des distributions limitées et, par conséquent, pour- breitung dieser Art. Drei der neuen Vorkommen scheinen raient être particulièrement sensibles à des per tur bations einen abgetrennten nördlichen Bestand in einer bio- locales, spécialement celles qui peuvent réduire le courant geografischen Provinz zu vertreten, wo die Art vorher comme des sécheresses prolongées et des activités hu maines unbekannt war. Das vierte Vorkommen dürfte als kleine comme le prélèvement d’eaux de surface ou souterraines. Erweiterung des bisher bekannten Bestandes einzuordnen sein. Die neu nachgewiesenen Vorkommen umfassen jeweils Sommario Habitate, wie sie ähnlich in den Gebieten des Kernbestandes In questo articolo viene documentata la presenza del pesce von C. rhombosomoides festgestellt worden waren; die arcobaleno di Cairns Cairnsichthys rhombosomoides Nichols Umweltdaten der neuen Verbreitungsgebiete werden doku- & Raven, 1928 in quattro località finora non censite. Le mentiert. Die Exemplare aus den nördlichen Vorkommen nuove località si trovano tutte al di fuori dell’area geografica unterscheiden sich genetisch, obwohl sie morphologisch den conosciuta per questa specie. Tre di queste sembrano rac - Tieren des Kerngebietes ähnlich erscheinen. Die neu fest- chiudere una popolazione settentrionale disgiunta posta in

155 aqua vol. 19 no. 3 - 19 July 2013 New Distribution Records for the Cairns Rainbowfish Cairnsichthys rhombosomoides: implications for conservation of a restricted northern population una provincia biogeografica in cui la specie era finora Four new locality records for C. rhombosomoides are sconosciuta. La quarta è un'estensione minore della popo- documented herein. The new localities are all out- lazione nota già esistente. Tutte le nuove località contengono side the currently recognised range for this species. habitat simili a quelli presenti nelle zone in cui sono diffuse Three of the new records appear to represent part of le popolazioni principali di C. rhombosomoides e per ciascuna di esse sono forniti i dati ambientali. Gli individui raccolti a disjunct northern population in a biogeographical nelle località settentrionali sono geneticamente distinti, province where the species was previously unknown. anche se morfologicamente simili a quelli della zona centrale The fourth record appears to be a minor range di distribuzione. Le popolazioni di C. rhombosomoides delle extension to the existing population. nuove località, avendo a disposizione un habitat molto ristretto e una limitata distribuzione, possono essere partico- METHODS larmente vulnerabili a perturbazioni locali, soprattutto The observations documented here form part of a quelle che possono ridurre la portata dei torrenti come una wider mapping project to better define the distribu- siccità prolungata o attività umane che comportano il prele- vamento di acque superficiali e sotterranee. tion of melanotaeniids across the Wet Tropics region. The project involves systematically searching for anomalous rainbowfish forms in known or sus- pected localities, based on published and unpub- INTRODUCTION lished reports, anecdotal evidence from local biolo- The Cairns rainbowfish (Cairnsichthys rhomboso- gists or aquarists, and the authors’ own experience in moides Nichols & Raven, 1928) is a small freshwater the region. fish (Atheriniformes: Melanotaeniidae) endemic to Collecting or observation methods were dependent Australia. When first described, it was placed within on local conditions. In streams where water clarity the genus Rhadinocentrus (Nichols & Raven 1928), was adequate, snorkel searching or bankside obser- but the taxon was later elevated to a new monotypic vations were used as a rapid assessment method. Fish genus Cairnsichthys (Allen 1980). were otherwise collected by dip net, particularly at Cairnsichthys rhombosomoides occurs only within night with the aid of spotlights. Data on water qual- the Wet Tropics bioregion (Sattler & Williams ity, environmental conditions and general fish com- 1999) of northeastern Queensland. Within that munities were collected at all sampling sites. bioregion, current knowledge indicates that it is fur- Generally, targeted streams were searched as far ther restricted to a few coastal drainages to the south upstream and downstream as was safe and practi- of Cairns. Originally, it was thought to occur only in cal. In some downstream areas, estuarine crocodiles the Russell-Mulgrave drainage system, but surveys were potentially present making it unsafe to sam- from the mid-1980s to early 2000s have extended ple, but these areas were generally unsuitable for C. the range southwards to include the North and rhombosomoides. Likewise, as soon as upstream South Johnstone rivers, Liverpool Creek, Maria areas became steep and torrential, it was considered Creek and the North Hull system (Kleiberg 1988; that this habitat was unsuitable and surveys were Pusey et al. 2004; Allen et al. 2002; Russell et al. concluded. When streams with C. rhombosomoides 1996). Thus, the currently recognised distribution of were located, an effort was made to radiate the sur- this species is confined to a coastal strip approxi- veys out to adjacent streams to determine the local mately 100 km long to the south of Cairns (Fig. 1). limits of distribution. The total area of this range is approximately 2,200 sq km, although much of this area has habitat unsuit- RESULTS able for C. rhombosomoides. This range correlates Three localities with anecdotal reports of C. rhom- very closely to the Wet Tropics Bioregional Province bosomoides were searched, including two localities in 3 (Innisfail) and the lower foothills of Province 7 the Daintree – Cape Tribulation area and one local- (Bellenden-Kerr-Lamb) (Sattler & Williams 1999). ity in the Yarrabah area. In the Daintree – Cape Within its known range, C. rhombosomoides has Tribulation area, the species was confirmed at one of very specific habitat requirements. It generally the localities, but could not be located at the other occurs in small well-shaded, clear permanent site. However, searches of other streams in the same streams below 100 m altitude, including lowland region resulted in the discovery of C. rhomboso- streams debouching into larger rivers, and upland moides at two additional localities. Cairnsichthys tributary streams in the foothills (Pusey & Kennard rhombosomoides was also confirmed to be still pre- 2004). sent at the Yarrabah locality. aqua vol. 19 no. 3 - 19 July 2013 156 Keith C. Martin and Susan Barclay

Fig. 1. New localities for C. rhombosomoides, and indicative limits of northern and southern populations.

157 aqua vol. 19 no. 3 - 19 July 2013 New Distribution Records for the Cairns Rainbowfish Cairnsichthys rhombosomoides: implications for conservation of a restricted northern population

Because the Daintree – Cape Tribulation localities about 600 m upstream of its confluence with the are clustered in an area that is significantly remote main part of Cooper Creek. Three C. rhomboso- from the previously known distribution limits of moides specimens were collected in the area up to this species (over 120 km to the north), C. rhombo- 100 m downstream of the road bridge, in pool and somoides from these locations are herein referred to riffle habitat. A search upstream of the road bridge as the “northern population” (Fig. 1). yielded more typical habitat and C. rhombosomoides Information on the occurrence of C. rhomboso- was found within a small stretch of watercourse up moides at the new localities (ordered from most to 500 m upstream of the road crossing. Investiga- northerly to most southerly) is provided in the fol- tions of another site some 2 km further upstream in lowing sections. The sections are divided into those the same tributary produced no C. rhombosomoides. from the “northern population” in the Daintree- Cooper Creek is a major coastal stream which Cape Tribulation region, and a site which is consid- debouches into the northern end of Alexandra Bay. ered an extension of the core or “southern popula- Little Cooper Creek is a small, second order tribution”. Results of fish community surveys conducted tary with a catchment size of about 1.5 km2, with simultaneously at the same sites are also presented. elevations from near sea level up to about 700 m. It drains steeply from the eastern side of Thornton Northern Population Sites Peak and joins Cooper Creek in the freshwater Little Cooper Creek reaches. Habitats along this creek are primarily Acting on anecdotal information (D. Sampson undisturbed rainforest, with some private residential pers. comm.), a locality in a tributary of Cooper and commercial properties. Creek (known locally as Little Cooper Creek) within At the locality where C. rhombosomoides was pre- the Daintree – Cape Tribulation area was searched sent, the creek flowed very clear, with pH of 7.5, by spotlighting on 10 May 2012. The locality was water conductivity (EC) of 23 µs and water temper- near a public road crossing (Turpentine Road), ature of 20°C. The creek had an average wetted

Fig. 2. C. rhombosomoides habitat, Little Cooper Creek. Photo by K. C. Martin. aqua vol. 19 no. 3 - 19 July 2013 158 Keith C. Martin and Susan Barclay width of 5 m, and depths of less than 0.5m over a Hutchinson Creek sandy substrate. The creek was 50% shaded by rain- On 10 May 2012 the authors conducted a walk- forest, with no aquatic vegetation. The C. rhomboso- ing/snorkelling visual survey of Hutchinson Creek, moides observation locality in Little Cooper Creek in the Daintree-Cape Tribulation area. The area was at S 16°10’10.75”; E 145°24’ 48.30” at an ele- from the Cape Tribulation road bridge up to a dis- vation of 22 m (Figs 1-2). tance of 1.5 km upstream was systematically South Cooper Creek searched by snorkelling and bankside visual assess- On 20 March 2013 the authors conducted a sur- ment over a period of 3.5 hours. At one point, a vey of a southern tributary of Cooper Creek within small log jam was encountered, and examination of the Daintree - Cape Tribulation area. There is some the pool immediately below indicated the presence confusion over the naming of this creek, and local of two sub-adult C. rhombosomoides. A single speci- property owners called it “Little Cooper Creek”, men was netted and retained. No other specimens of which is the same name regularly given to the previ- C. rhombosomoides were located during the survey. ous site. To avoid confusion with the latter site, we Other tributaries of Hutchinson Creek with appar- refer to this locality as South Cooper Creek. ently suitable habitat, including Fairy, McLean and A transect of approximately 300 m along this creek Buchanan Creeks were also searched, but no C. either side of the main Cape Tribulation road was rhombosomoides were observed. searched by snorkelling in the daytime and by spot- Hutchinson Creek is a moderately sized third order lighting at night. In a shallow, fast flowing section of stream, which has its headwaters in the Thornton the creek immediately downstream of the road, a Range and debouches into the southern section of group of five C. rhombosomoides were observed by Alexandra Bay. It is the next major catchment south snorkelling. Night spotlighting in the same area and of Cooper Creek. The freshwater catchment size is further upstream yielded two additional specimens. about 18 km2, with elevations from near sea level up South Cooper Creek is a moderately sized, peren- to about 300 m. There is a significant estuarine sec- nial third order stream, which has its headwaters in tion, generally commencing downstream of the the Thornton Range and debouches into the main Cape Tribulation road crossing. The freshwater Cooper Creek estuary at the northern end of Alexan- reaches include foothill habitats within Cape Tribu- der Bay. Although this creek is a tributary of Cooper lation National Park, and lowlands running through Creek, the two watercourses do not join until the privately owned horticultural lands. estuarine section of the complex is reached. Further, At the time of the surveys, the creek flowed very anecdotal evidence from local property owners indi- clear, with pH of 7.1, water conductivity (EC) of 38 cates that the lower reaches of this creek flow under- µs and water temperature of 23°C. The creek in the ground – at least during drier parts of the year. surveyed area was about 10 m wide, with average The freshwater catchment size is about 5 km2, with depths of about 1 m and some very deep pools (up elevations from near sea level up to about 700 m. to 4 m deep). Substrates were primarily sand or The creek runs through primarily undisturbed rain- coarse gravel. The creek was shaded about 50%, pre- forest for most of its length, although the midreach dominantly by fringing rainforest, although some sections where C. rhombosomoides occurs are within bankside areas had been cleared. Aquatic vegetation private allotments. At the time of the surveys, the was absent or very sparse in most areas. The C. creek flowed very clear, with pH of 6.5, water con- rhombosomoides observation locality in Hutchinson ductivity (EC) of 27 µs and water temperature of Creek was at S 16°12’59.23”; E 145°24’55.45” at an 24.5°C. The creek in the surveyed area was about 10 elevation of 24 m (Figs 1&3). m wide, with average depths of about 1 m and max- Forest Creek Tributary imum depths of 2 m, although in the furthest down- A single specimen of a fish identified in the field as stream reaches, large pools of over 4 m depth were C. rhombosomoides was collected from a small tribu- observed. Substrates were primarily fine or coarse tary of Forest Creek (itself a northern tributary of the gravel. The creek was shaded about 50%, predomi- Daintree River) during the course of a Government nantly by fringing rainforest. Aquatic vegetation was fish survey in 2004 (F. Kroon, pers. comm). Confir- absent or very sparse in most areas. The C. rhombo- mation of this identification could not be obtained. somoides observation locality in South Cooper Creek This locality, about 4 km south of the Hutchinson was at S 16°10’ 35.65”; E 145°24’57.39” at an ele- Creek site, but in a different catchment, was snorkel vation of 25 m. searched by the authors on 7 March 2013 through-

159 aqua vol. 19 no. 3 - 19 July 2013 New Distribution Records for the Cairns Rainbowfish Cairnsichthys rhombosomoides: implications for conservation of a restricted northern population

Table I. Fish community composition at four new localities where C. rhombosomoides occurs.

Common Name Scientific Name Hutchinson Little Cooper South Cooper Yarrabah Ck Ck Ck Long-finned Eel Anguilla reinhardti xxx Yellow-finned Tandan Neosilurus hyrtlii x Northern Cobbler Tandanus cf. tandanus xx x Cairns Rainbowfish Cairnsichthys rhombosomoides xx xx Eastern rainbowfish Melanotaenia s. splendida xx xx Banded Rainbowfish Melanotaenia trifasciata xx Pacific Blue-eye Pseudomugil signifer xx x Sail-fin Glassfish Ambassis agassizii x Glassfish Ambassis miops x Margined Flagtail Kuhlia marginata x Jungle Perch Kuhlia rupestris xx x False Celebes Goby Glossogobius illimus x Silver Grunter Mesopristes argenteus x Roman Nosed Goby Awaous acritosus xx Empire Gudgeon Hypseleotris compressa xx xx Snakehead Gudgeon Giurus margaritacea xx Green-backed Gauvina Bunaka gyrinoides xx x Mangrove Jack Lutjanus argentimaculatus x out its entire freshwater length of suitable habitat sent, the creek flowed very clear, with pH of 7.6, (about 500 m). Although there was excellent C. water conductivity (EC) of 68 µs and water temper- rhombosomoides habitat in this creek, no specimens ature of 23°C. The creek had an average width of were located. The 2004 record from the Forest Creek 1.5 m, and depths of less than 0.3 m over a sandy tributary was at S 16°14’45.7”; E 145°23’35” at an substrate. The creek was 70% shaded by rainforest elevation of 15 m. This and four other tributaries of with no aquatic vegetation. The Yarrabah observa- Forest Creek were searched by the authors on several tion was at S 16°55’28.36”; E 145°54’12.23” at an occasions in 2012 and 2013, but C. rhombosomoides elevation of less than 10 m (Fig. 1). was not located despite suitable habitat. Fish Communities Southern Population Site A total of 18 fish species including C. rhomboso- Unnamed Creek, Yarrabah moides were recorded at the four new localities dur- Acting on anecdotal information (G. Aland, pers. ing the course of the surveys. These included 14 in comm.), a locality on the Yarrabah Peninsula to the Hutchinson Creek, 11 in Little Cooper Creek, nine east of Cairns City was inspected on 17 May 2012 in South Cooper Creek and six at the Yarrabah local- and a reported population of C. rhombosomoides was ity (Table I). Species such as: Melanotaenia s. splen- found there. A few other sites in the general area of dida Peters, 1866; Tandanus cf. tandanus Mitchell, Yarrabah were also inspected by visual bankside 1838; Kuhlia rupestris Lacepède, 1802; Pseudomugil observations or snorkel survey, but no other C. signifer Kner, 1866; and Hypseleotris compressa Krefft, rhombosomoides localities were found. 1864 were generally common at all or most of the The Yarrabah locality is a small, second order sites. Cairnsichthys rhombosomoides occurs along with stream which flows down the western slope of Grant two other rainbowfish species (Melanotaenia s. splen- Hill, to disperse into a large freshwater swamp that dida and M. trifasciata Rendahl, 1922) in the two eventually debouches into Mission Bay. The Cooper Creek tributaries. The occurrence of M. tri- upstream catchment size of this stream is extremely fasciata in the Cooper Creek system has been previ- small, about 0.3 sq km, consisting primarily of ously noted (Ebner & Thuesen 2010; Martin 2010) undisturbed rainforest, with elevations from near sea and appears to be an isolated population which is level to 200 m. probably the most southerly locality for this species At the locality where C. rhombosomoides was pre- on the east coast of Australia. In this area, the distri- aqua vol. 19 no. 3 - 19 July 2013 160 Keith C. Martin and Susan Barclay butions of C. rhomosomoides and M. trifasciata over- species elsewhere to date. Other creeks which have lap, probably the only location in Australia where been investigated include: additional tributaries of sympatry between these two species occurs. Cooper and Hutchinson creeks; Forest Creek and Anecdotal evidence from local aquarists suggests other small tributaries of the Daintree River imme- that a further rainbowfish species, Melanotaenia diately to the south of Hutchinson Creek; the maccullochi Ogilby, 1915 may also be present in the Noah/Oliver and Myall Creek systems immediately Cooper Creek catchment. If so, and given the small to the north of Cooper Creek; and Emmagen Creek size of the catchment, this would make Cooper to the north of Cape Tribulation. Creek one of the richest sites for Melanotaeniidae in In addition to the authors’ own surveys, we note Australia. Paradoxically, fish communities of the that other intensive fish surveys covering almost all Daintree – Cape Tribulation streams have been major streams in the Daintree – Cape Tribulation shown to have a close affinity to high island streams area have been conducted over the past few years of the Pacific islands where communities are domi- and that C. rhombosomoides has not been recorded nated by amphidromous species (Thuesen et al. elsewhere in the region to date (B. Ebner; P. Thue- 2011). That Cooper Creek may also have a very sen, pers. comm.) high diversity of Melanotaeniidae (a primary fresh- The discovery of C. rhombosomoides at three locali- water fish group) which are geographically isolated ties in the Daintree – Cape Tribulation region (and adds another dimension to the conservation and the likelihood that it occurs, or previously occurred ecological values of this area. at a fourth locality) extends the range of this species some 120 km to the north of the currently recog- DISCUSSION nised range and into a biogeographical province (9 – Searches for Cairnsichthys rhombosomoides at other Daintree – Bloomfield) where it was previously localities in the Daintree – Cape Tribulation region unknown. using the same methodology (primarily snorkel In the Cooper Creek system, C. rhombosomoides is searching or spotlighting) have failed to locate the now known from two disconnected tributaries, but

Fig. 3. C. rhombosomoides collection site, Hutchinson Creek. Photo by K. C. Martin

161 aqua vol. 19 no. 3 - 19 July 2013 New Distribution Records for the Cairns Rainbowfish Cairnsichthys rhombosomoides: implications for conservation of a restricted northern population the status of this species in other parts of the Cooper two individuals were observed and a thorough inves- Creek catchment is unclear. In Little Cooper Creek tigation of a significant length of the same stream it is restricted to a lower reach area of less than 1 km failed to locate further specimens. It is postulated in length. It is not present in the upper reaches of that the specimens observed at this location are this tributary above a series of rapids, nor does it likely to represent straying individuals from an as yet appear to be present in the main channel of Cooper undiscovered core population, probably located in a Creek itself. In a significant study of cling-gobies in small tributary of Hutchinson Creek further the “Blue Hole” area of Cooper Creek (approxi- upstream. This stream has been surveyed only a few mately 500 m upstream of the confluence with Lit- hundred metres further upstream of the observation tle Cooper Creek), 19 species of freshwater fishes site, so the status of C. rhombosomoides in the upper were recorded, but C. rhombosomoides was not one tributaries is still unknown. of them (Ebner & Thuesen 2010). The confirmation of isolated C. rhombosomoides In South Cooper Creek, there is only a small area populations in the Daintree – Cape Tribulation of suitable habitat downstream of the main Cape streams remote from the core population base of this Tribulation road crossing, then the creek becomes species to the south of Cairns raises questions as to very large and deep as it reaches the estuary. The area whether the newly discovered northern populations upstream of the road crossing with suitable habitat may be distinctive, may have been translocated, or if has not yet been determined, but is likely to be less further undiscovered populations may exist in the than 2 km in length. intervening area or even in areas further north. The Hutchinson Creek site, also in the Daintree – To assess the distinctiveness of these fish from the Cape Tribulation region, is unusual in that C. rhom- southern population, representative samples were bosomoides was observed in the main channel of a collected from Little Cooper Creek (series of ten larger, more open creek than at the other sites. Only individuals) and Hutchinson Creek (a single indi-

Fig. 4. Cairnsichthys rhombosomoides “northern population” from Little Cooper Creek. Photo by K. C. Martin

Fig. 5. Cairnsichthys rhombosomoides “southern population” from Harvey Creek. Photo by K. C. Martin. aqua vol. 19 no. 3 - 19 July 2013 162 Keith C. Martin and Susan Barclay vidual) and forwarded live to the Northern Territory extent. It generally occurred only in a small, perma- Museum for comparison with specimens from loca- nently flowing section of the stream between the tions in the Russell-Mulgrave and Johnstone River base of the mountain range and the downstream sec- catchments within the normal range of this species. tions where depths are increased and flow is This research is currently underway, although pre- reduced. Within that range, the species was further liminary results indicate that specimens from the confined to a microhabitat of shallow, sandy and northern population are genetically dissimilar to swiftly flowing sections which are heavily shaded those from southern populations, therefore dis- and contain refuge sites such as woody debris, counting the possibility of translocation. Superfi- undercut banks and instream vegetation such as ter- cially, live C. rhombosomoides from Cooper and restrial tree roots. There is an abrupt downstream Hutchinson creeks appear similar to fish from the cessation of C. rhombosomoides populations where southern populations. The most obvious difference the creeks transition from these habitats into deeper is the northern population fish have pronounced pools and more open, slow flowing runs. These blue colouration in their eyes and to a lesser extent, downstream habitats not only are seemingly unsuit- around the gills and upper body (Fig. 4). These fea- able for C. rhombosomoides, but also contain more tures can be seen in some examples from the south- predator species such as Lutjanus argentimaculatus ern populations, but they are far more pronounced Forsskål, 1775. Thus, it seems unlikely that C. rho- and brighter in the Cooper-Hutchinson fish. These mosomoides could migrate between creek systems via features are not evident in the Yarrabah population, downstream movements and therefore would be which closely resembles specimens from the south- unable to populate other areas by this method. In ern populations and is also genetically similar to the case of Cooper Creek, the two known popula- samples from the southern population. A compara- tions are quite close to each other (1 km overland) tive example of a C. rhombosomoides from Harvey but the streams are separated downstream by a Creek in the Russell River catchment and part of the seemingly impenetrable barrier of deep pool and southern population, is shown in Fig. 5. estuarine habitat. The possible occurrence of C. rhombosomoides in Thuesen et al. (2008) found that most C. rhombo- small streams draining into Trinity Inlet east of somoides populations at sites within the southern Cairns (i.e. the north-western side of the Yarrabah population were typically isolated, both physically Peninsula) has been suggested by Pusey et al. (2004), and genetically, and may also be fragmented due to since this area may have historical connections to the predator pressure in downstream areas. It appears Mulgrave River. Cairnsichthys rhombosomoides has that the northern population sites have the same not yet been found in these streams, although access characteristics. These small, isolated populations to suitable habitat sites is difficult. The Yarrabah site may be particularly vulnerable to localised extinc- is in the same general area, but on the eastern side of tion. the Murray Prior Range. The failure to locate C. rhombosomoides in the For- The likelihood of C. rhombosomoides occurring in est Creek area may be significant. It is considered the intervening coastal streams between Cairns and very likely that the original observation is genuine, the Daintree River is considered low. Most of these since the specimen was collected and identified by a streams are steep and short, and generally do not professional ichthyologist, and the habitat is ideal. contain optimal habitat. Also, these streams have Our experience with snorkel searching in these clear been surveyed both by professional biologists (e.g. creeks is that although C. rhombosomoides is gener- Pusey & Kennard 1994; Russell et al. 1998; Russell ally uncommon, if it is present in an area it can usu- et al. 2000) and by knowledgeable amateur aquar- ally be located. Our extensive surveys of the Forest ists, and this species has not been reported to date. Creek tributaries have failed to locate the species Apparently suitable habitat occurs in streams to the there and it is possible that the species may have dis- north of Cooper Creek, in the Cape Tribulation and appeared from these creeks over the past ten years. If Bloomfield regions, although to date there is no that is the case, then the future of the remaining new indication of this species presence in these areas populations is also insecure. despite significant survey effort. Cairnsichthys rhombosomoides is not currently listed All of the new localities can be regarded as foothill under any Australian legislation as a threatened tributary habitats. At every locality, the species was species, but is listed on the IUCN’s Red Data List as found to be uncommon and highly restricted in ‘Vulnerable” (Wager 1996). The small extent of

163 aqua vol. 19 no. 3 - 19 July 2013 New Distribution Records for the Cairns Rainbowfish Cairnsichthys rhombosomoides: implications for conservation of a restricted northern population these populations and the shallow water environ- REFERENCES ments in which they occur make them extremely ALLEN, G. R. 1980. A generic classification of the rainbow- susceptible to any localised disturbances. Extreme fishes (family Melanotaeniidae). Records of the Western Aus- weather conditions, particularly extended drought, tralian Museum 8: 449-90. could have a severe impact on these populations if ALLEN, G. R., MIDGLEY, S. H. & ALLEN, M. 2002. Field Guide to the Freshwater Fishes of Australia. Western Aus- the small streams they inhabit stopped flowing or tralian Museum: Perth. 394 pp. dried completely. This could even lead to a perma- EBNER, B. C. & THUESEN, P. 2010. Discovery of stream- nent local extinction. Similarly, human activities cling-goby assemblages (Stiphodon species) in the Aus- such as localised habitat clearing, ground and sur- tralian Wet Tropics. Australian Journal of Zoology 58: 331- face water extraction and stream pollution could 340. have severe and possibly extreme effects on these KLEIBERG, T. 1988. The Cairns Rainbowfish Cairnsichthys populations. rhombosomoides (Nichols & Raven). Fishes of Sahul 5 (2): Based on current knowledge, it must be assumed 215-216. MARTIN, K. C. 2010. Along the Bloomfield Track, North that the northern population of this species is vul- Queensland. In-Stream, 19 (10). nerable to extinction and therefore requires urgent NICHOLS, J. T. & RAVEN, H. C. 1928. A New Melano- conservation actions. These measures may include: taeniin fish from Queensland. American Museum Novitates formal steps to have the form listed under State or 296. American Museum of Natural History. Commonwealth threatened species legislation; alert- PUSEY, B. J., KENNARD, M. J. & ARTHINGHTON, A. H. ing appropriate State or Local authorities (including 2004. Freshwater Fishes of North-eastern Australia. CSIRO managers of the nearby National Park) to the pres- Publishing: Melbourne. 684 pp. ence of this fish in the region; and ensuring that RUSSELL, D. J., HALES, P. W. & HELMKE, S. A. 1996. Stream Habitat and Fish Resources in the Russell and Mulgrave future approvals for actions such as planning, devel- Rivers Catchments. Queensland Department of Primary opment, vegetation removal, water extraction or Industries, Brisbane. 68 pp. other potential habitat disturbances in the area RUSSELL, D. J., MCDOUGALL, A. J. & KISTLE, S. E. 1998. should take the protection of this species into con- Fish Resources and Stream Habitat of the Daintree, Saltwa- sideration. ter, Mossman and Mowbray Catchments. Information Series Further research into the taxonomic status and dis- QI98062. Queensland Department of Primary Industries, tribution extent of the C. rhombosomoides form in Brisbane. 72 pp. the Daintree – Cape Tribulation region is recom- RUSSELL, D. J., MCDOUGALL, A. J., RYAN, T. J., KISTLE, S. E., ALAND, G., COGLE, A. L. & LANGFORD, P. A. 2000. mended, and the status of existing populations Natural Resources of the Barron River Catchment 1. Stream should be monitored. Additionally, it may be useful habitat, fisheries resources and biological indicators. Informa- to ensure that captive breeding populations be main- tion Series QI00032. Queensland Department of Primary tained as a security measure against local extinctions. Industries, Brisbane. 108 pp. SATTLER, P. & WILLIAMS, R. (Eds.) 1999. The Conservation ACKNOWLEDGEMENTS Status of Queensland’s Bioregional Ecosystems. Published The authors wish to acknowledge Mr. Dean Samp- by Environmental Protection Agency, Brisbane. son of Mossman, Qld, and Mr. Glynn Aland of THUESEN, P. A., PUSEY, B. J., PECK, D. R., PEARSON, R. & CONGDON, B. C. 2008. Genetic differentiation over small Beaudesert, Qld for alerting us to the possible occur- spatial scales in the absence of physical barriers in an Aus- rence of C. rhombosomoides at Little Cooper Creek tralian rainforest stream fish. Journal of Fish Biology 72 (5): and Yarrabah, respectively. Dr. Frederieke Kroon 1174-1187. provided information on a fish survey at Forest THUESEN, P. A., EBNER, B. C., LARSON, H., KEITH, P. & Creek. Dr. Michael Hammer provided advice and SILCOCK, R. M. 2011. Amphidromy Links a Newly Doc- encouragement, and arranged for specimens to be umented Fish Community of Continental Australian lodged at the Northern Territory Museum. Dr. Streams, to Oceanic Islands of the West Pacific. PLoS ONE Brendan Ebner provided institutional support for 6 (10): e26685.doi:10.1371/journal.pone.0026685 WAGER, R. 1996. Cairnsichthys rhombosomoides. In: IUCN permitting, and also provided information on results 2012. IUCN Red List of Threatened Species. Version of recent fish research in the Daintree – Cape Tribu- 2012.2. . Downloaded on 28 Octo- lation area. Dr. Michael Hammer and Dr. Gerald ber 2012. Allen kindly reviewed an early draft of the paper. This research was conducted under James Cook University Animal Ethics Approval No. A1873 and Qld General Fisheries Permit 162324. aqua vol. 19 no. 3 - 19 July 2013 164 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.): low the format set out in previous publications of aqua. 197-221. 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): species are followed by the systematic group to which they 72-79. 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. 19 (3), 19 July 2013

Contents:

Dalton Tavares Bressane Nielsen and Roger Brousseau: Description of a new species of annual fish of the genus Neofundulus (Cyprinodontiformes: Rivulidae) from the upper río Mamoré basin, Bolivia ...... 109-114

Dalton Tavares Bressane Nielsen and Roger Brousseau: Spectrolebias pilleti, a new annual Killifish (Cyprinodontiformes: Rivulidae: Cynolebiatinae) from the upper río Mamoré basin, Bolivia ...... 115-122

Gerald R. Allen, Mark V. Erdmann and Christine L. Dudgeon: Hemiscyllium halmahera, a new species of Bamboo Shark (Hemiscylliidae) from Indonesia ...... 123-136

Gerald R. Allen and Renny K. Hadiaty: Melanotaenia sneideri, a New Species of Rainbowfish (Melanotaeniidae), from West Papua Province, Indonesia ...... 137-146

Helen K. Larson: A new species of Egglestonichthys from northern Australian estuaries (Teleostei, Gobiidae, Gobiinae) ...... 147-154

Keith C. Martin and Susan Barclay: New Distribution Records for the Cairns Rainbowfish Cairnsichthys rhombosomoides (Melanotaeniidae): implications for conservation of a restricted northern population ...... 155-164

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: Neofundulus splendidus, male, holotype, 63.5 mm SL: Bolivia, Departamento de Santa Cruz, San Miguel. Photo by G. Dethu.

New endemic Midas cichlid species from Niacaraguan lake to be described in a upcoming issue of aqua.