aqua Journal of Ichthyology and Aquatic Biology Vol. 11 (1), February 2006

Aquapress ISSN 0945-9871 aqua - Journal of Ichthyology and Aquatic Biology

Managing Editor: Scope and aims is an international journal which publishes original Heiko Bleher aqua scientific articles in the fields of systematics, taxonomy, Via G. Falcone 11, 27010 Miradolo Terme (PV), Italy biogeography, ethology, ecology, and general biology of Tel.: +39 0382 754707/08 - Fax: +39 0382 754129 fishes, amphibians, aquatic invertebrates, and plants. e-mail: [email protected] Papers on freshwater, brackish, and marine organisms will be considered. aqua is fully refereed and aims at Scientific Editor: publishing manuscripts within 2-4 months of acceptance. With the publication of aqua we are pursuing a new con - Dr. Walter Ivantsoff cept: In view of the importance of colour patterns in Senior Research Fellow, species identification and animal ethology, authors are Department of Biological Sciences, encouraged to submit colour illustrations as well as Macquarie University, N.S.W. 2109, Australia descriptions of coloration. It is our aim to provide Tel. +61 2 9850 8167 - Fax +61 2 9869 8886 the international scientific community with an efficiently e-mail: [email protected] published series meeting high scientific and technical standards. Editorial Board: Call for papers Gerald R. Allen, I Dreyer Road Roleystone, W. A. Australia 6111 The editors welcome the submission of original manu - scripts which should be sent directly to the scientific editor. George W. Barlow, Department of Integrative Biology, Full length research papers and short notes will be consid - University of California, Berkeley, CA 94720-3140, ered for publication. There are no page charges and colour U.S.A. illustrations will be published free of charge. Authors will receive 1 free reprint and 1 PDF-file of each paper. Henri J. Dumont, Rijksuniversiteit Gent, Laboratorium voor Ecologie der Dieren, Zoogeografie en Natuur- behoud, K. L. Ledeganckstraat, 9000 Gent, Belgium Subscription Notice A volume (4 issues) of will be published each year, Jacques Géry, Chemin du Plantier, 24200 Sarlat, aqua France each issue comprising 48 pages (including cover). The annual subscription rate (for one volume = 4 issues) Frank Kirschbaum, Institut für Gewässerökologie und is Euro 40.00 (US$ 48.00) plus postage Euro 10.00 Binnenfischerei, Abt. 4 Forschungsverbund Berlin e. V. (US$ 12.00) and for priority postage Euro 17,00 Müggelseedamm 310, 12587 Berlin, Germany (US$ 20.00). Subscription enquires should be sent to the address Friedhelm Krupp, Forschungsinstitut Senckenberg, given below or our e-mail: [email protected] Senckenberganlage 25, 60325 Frankfurt am Main, Germany aqua binder Christian Lévêque, CNRS - Programme Environnement Binders for Volumes of Journal Ichthyology and Vie et Sociétès, 1 Place Aristide Briand, 92195 Paris aqua Aquatic Biology are available at cost price Euro 12,50 Cédex, France (US$ 15.00) plus postage Euro 8,00 (US$ 10.00). Volker Mahnert, Muséum d’Histoire Naturelle, Route de Notice: aqua Volumes 1(1)-5(4) = 1. binder; Malagnou 1, 1211 Genève 6, Switzerland Volumes 6(1)-9(4) = 2. binder. Paolo Parenti, Department of Enviromental Sciences, University of Milan-Bicocca, Piazza della Scienza 1, Special Publication I-20126 Milan, Italy From 2003 onwards Aquapress will publish a series of Special Publications . The first Special Publication of John E. Randall, Bishop Museum, 1525 Bernice Street, aqua will be available in December 2003. From then on, P.O. Box 19000-A, Honolulu, Hawaii, U.S.A. Special Publications will be printed yearly, or more often. All Special Publications will contain around 100 pages or Wolfgang Schneider, Hessisches Landesmuseum, more and will only be available separately from normal Darmstadt, Friedensplatz 1, 64283 Darmstadt, issues of aqua . Enquiries about subscriptions and the Germany prices of Special Publications should be sent to the address given below or via e-mail: [email protected] Mário de Pinna, Museu de Zoologia da USP, Av. Nazaré 481, São Paulo-SP 04263-000, Brazil Wolfgang Villwock, Universität Hamburg, Zoologisches ISSN 0945-9871 Institut und Zoologisches Museum, Martin-Luther-King- Publisher: Aquapress, Redazione aqua, Platz 3, 20146 Hamburg, Germany I-27010 Miradolo Terme (Pavia), Italy Printer: Grafiche Dessì s.r.l. (Torino), Italy Chem Yi-yu, Institute of Hydrobiology, Academia Sinica, Typesetting: Rossella Bulla Wuhan Hubei, P. R. China © 2006 aqua , Journal of Ichthyology and Aquatic Biology aqua, Journal of Ichthyology and Aquatic Biology

Cirrhilabrus brunneus , a new wrasse (Pisces: Labridae) from north-eastern Kalimantan, Indonesia

Gerald R. Allen

Department of Aquatic Zoology, Western Australian Museum, Francis Street, Perth, WA 6000, Australia

Accepted: 09.02.2006

Keywords Materials and Methods Taxonomy, marine fishes, Cirrhilabrus , new species, Counts of fin spines are given in Roman numerals and Labridae, Kalimantan, Indonesia soft rays in Arabic. Pectoral ray counts include the rudi - mentary upper ray. The lateral line is interrupted; the Abstract count of the anterior part is given first, followed by a plus Cirrhilabrus brunneus is described from a single male sign and the peduncular part. Only lateral line scales specimen, 43.6 mm SL, collected at north-eastern Kali - with tubes are counted. All the tubed scales of the mantan, Indonesia. It is one of only three species in the peduncular part are counted, even though one is usually genus that possesses a lunate caudal fin. The new located posteriorly to the base of the caudal fin. The species, which is overall dark brown in colour, including number of scales in the rows on the cheek is counted the median and pelvic fins, closely resembles C. lunatus from where they commence below the front of the orbit from the Ryukyu Islands, southern Japan, and the Oga - to behind the centre of the orbit. Gill raker counts include sawara Islands. However, unlike the latter species, it all rudiments. Because it may be difficult to determine lacks a broad orange-yellow zone along the sides at the which raker is at the angle, only the total gill raker count level of the pectoral fins, a pale yellowish band along the is given. base of the dorsal fin, and irregular diagonal red lines on Lengths of specimens are given as standard length the snout and the postorbital portion of the head that (SL) except estimates of total length (TL) of fishes pho - extends onto the anterodorsal part of the body. There is tographed underwater; this is the straight-line measure - also a substantial difference in maximum size with ment from the front of the upper lip to the base of the males of C. lunatus attaining a maximum standard caudal fin (end of hypural plate). Measurements are length of about 85 mm in comparison with less than 50 given as percentages of the standard length. Head mm for C. brunneus. length is the distance from the front of the upper lip to the posterior end of the opercular membrane. Body depth is Introduction the greatest depth to the base of the dorsal fin (adjust - The labrid genus Cirrhilabrus Temminck and Schlegel ing for any malformation of the abdomen due to preser - contains small, colourful, sexually dimorphic fishes that vation). Body width is measured just posterior to the inhabit Indo-west Pacific coral reefs. Allen (2000) listed opercular flap. Snout length is taken from the front of the 36 species, but since then six additional species have upper lip to the fleshy edge of the orbit (if the upper jaw been described by Senou and Hirata (2000), Randall is protruded, it is pressed back to the non-protractile and Pyle (2001), Randall and Nagareda (2002), and position before the measurement is taken. The same is Allen et al . (2003). true of SL and head length measurements. Orbit diam - The members of the genus typically live in rubble habi - eter is the greatest fleshy diameter. Interorbital width is tats, frequently below 20 m, and feed on zooplankton the least bony width. Caudal peduncle depth is the least well above the bottom. Due to their relatively deep- depth; caudal peduncle length is the horizontal mea - dwelling habits, most species remained undetected until surement between verticals at the rear base of the anal the widespread use of scuba equipment by scientific fin and the caudal fin base. Measurements of fin spines divers over the past few decades. No doubt additional and rays are taken to the extreme base of these ele - discoveries will occur in the future, especially with the ments. Caudal concavity is the horizontal distance increased use of specialized deep diving equipment between the tips of the longest and shortest caudal rays. such as mixed-gas scuba and small research sub - Pectoral fin length is taken from the tip of the longest ray marines. to the base of this ray. Pelvic fin length is measured from The present paper describes a new species that was the base of the spine to the tip of the longest ray. collected by the author during a recent coral reef survey The holotype and only known specimen is deposited for The Nature Conservancy at the Berau district of at Pusat Penelitian dan Pengembangan Oseanologi, north-eastern Kalimantan, Indonesia. Jakarta, Indonesia (NCIP).

1 aqua vol. 11 no. 1 - 2006 Cirrhilabrus brunneus , a new wrasse (Pisces: Labridae) from north-eastern Kalimantan, Indonesia

Cirrhilabrus brunneus n. sp. pointed, its length 3.9 (8.7) in head; orbit diameter 3.4 Dusky Wrasse (Fig. 1) (9.9) in head; interorbital space slightly convex medially, strongly convex laterally, the least bony width 4.4 (7.8) Holotype: NCIP 6306, male, 43.6 mm, Kaniungan in head; caudal peduncle depth 2.3 (14.7) in head; cau - Besar Island, East Kalimantan, Indonesia (01°06.934’N, dal peduncle length 1.9 (17.7) in head. 118°50.127’E), 40 m, collected with quinaldine sulphate Mouth terminal and oblique, forming an angle of about and hand net by G. Allen, 20 October 2003. 30 degrees to horizontal axis of body; mouth small, the maxilla just reaching a vertical at posterior nostril, the Diagnosis upper jaw length 4.2 (8.0) in head; dentition of holotype Dorsal rays XI,9; anal rays III,9; pectoral rays 15; lat - typical of the genus with three pairs of canine teeth eral line scales 17 + 6; median predorsal scales 5; hor - anteriorly at side of upper jaw, the first forward project - izontal scale rows on cheek below eye 2; gill rakers 13; ing, the next two strongly recurved and outcurved, the body depth 3.1 in SL; head length 2.9 in SL; snout third much the longest; an irregular row of very small length 3.9 in head; lunate caudal fin of male with pro - conical teeth medial to upper canines; side of upper jaw duced lobes; pelvic fins relatively short, not reaching with about 16 small conical teeth; lower jaw with a sin - anal fin origin when depressed; diagnostic live colour gle pair of forward projecting canines and a row of very pattern features of males include a dusky brown overall small conical teeth in the symphyseal gap; side of lower coloration with bronze hue on breast and belly, dark jaw with a row of about 22 small conical teeth, decreas - brown median and pelvic fins with broad white posterior ing in size posteriorly. Gill rakers small, the longest on margins on dorsal and caudal fins, and dorsal and ven - first branchial arch less than half length of longest gill fil - tral edges of caudal fin with prominent brown margins aments. that taper to a point posteriorly. Posterior margin of preopercle with 26 fine serrae; margins of posterior and ventral edges of preopercle Description free to about level of middle of pupil. Anterior nostril Proportional measurements expressed in thousandths small and inconspicuous, in a short membranous tube of the standard length are provided in parenthesis in the with a posterior flap, located anterior to upper edge of following paragraphs. eye nearly one half distance to front of upper lip; aper - Dorsal rays XI,9; anal rays III,9; dorsal and anal soft ture of posterior nostril much larger than any head rays branched except first and second rays of dorsal fin pores, without elevated rim, located posterior and and first ray of anal fin; the last dorsal and anal soft rays slightly dorsal to anterior nostril on a vertical with ante - branched to base; pectoral rays 15, the upper two rior, bony edge of orbit. Pores of cephalic lateralis sys - unbranched; pelvic rays I,5; principal caudal rays 13, tem adjacent to ventroposterior half of orbit 15; a series the upper and lower rays unbranched; upper and lower of 9 pores along margin of preopercle linking with 4 on procurrent caudal rays 4; lateral line 17 + 6; scales mandible to front of chin; a series of 12 pores from above lateral line to origin of dorsal fin 2; scales below above upper edge of preopercle passing dorsal to orbit lateral line to anus 6; median predorsal scales 5; and ending in front of anterior nostril; a series of 9 pores median preventral scales 5; transverse scale rows on on each side of head from first lateral line scale to front cheek 2, the upper row with 6 and the lower row with 9 of scaled part of nape, plus 3 mid-interorbital pores. scales; circumpeduncular scales 15; gill rakers 13. Scales cycloid; head scaled except snout, interorbital Body moderately elongate, the depth 3.1 (32.1) in SL; space, and ventrally; lowermost of two transverse rows body compressed, the width 1.9 (16.7) in depth; dorsal of scales below eye larger than upper; naked flange of profile of head nearly straight, becoming slightly convex ventral edge of preopercle about half height of lower row on nape; head length 2.9 (33.9) in SL; snout moderately of scales; base of dorsal and anal fins with a row of large

Fig. 1. Cirrhilabrus brunneus n. sp. , freshly collected male holotype, 43.6 mm SL, Fiji Islands. Photo by G. R. Allen. aqua vol. 11 no. 1 - 2006 2 Gerald R. Allen elongate scales, one per membrane (except first scale Colour or male holotype in alcohol: after more than which covers membranes of first and second spines), two years of preservation, the colour pattern is remark - the longest about one-half spine length (basal scales ably similar to that described in the previous paragraph progressively shorter posteriorly on membranes of soft with the exception of a tan iris and tannish lower one- portion of fin); peduncular lateral line scales followed by third of the body. one slightly larger pored scale (included in lateral line Live colour of female from underwater field count) on base of caudal fin with a slightly posterior notes: overall brownish (red when illuminated with scale above and below, these three scales followed by underwater torchlight) with faint longitudinal dark a vertical row of three large scales, the middle one over - stripes on head and body and small dark spot on lapping the ones above and below, reaching two-thirds upper caudal peduncle. distance to posterior margin of fin; pectoral fins scale - less; pelvic fins with a median ventral process of two Remarks elongate scales, the more pointed posterior scale The caudal fin shape of the male serves to separate C. extending to tip of depressed pelvic spine spine; slender brunneus from nearly all other members of the genus axillary scale of each pelvic fin extending about equal in which generally have truncate, emarginate, rhomboid, length to pelvic spine. or lanceolate caudal fins. The only other species with Origin of dorsal fin above third lateral line scale; first produced fin lobes are C. lunatus Randall and Masuda dorsal spine 4.8 (7.1) in head; remaining dorsal spines (1991) from the Ryukyu Islands, southern Japan, and progressively longer, the last 1.9 (18.1) in head; inter - the Ogasawara Islands, and C. johnsoni Randall (1988) spinous membranes of dorsal fin extending above spine from the Marshall Islands. Males of both species have tips, supported by the terminal cirrus projecting upward strongly lunate caudal fins, with the upper and lower from just behind each spine tip; first to third dorsal soft rays forming filamentous extensions. rays the longest, 2.1 in head; origin of anal fin on a ver - The distinctive colour patterns of the three species tical with base of penultimate dorsal spine; first anal with lunate-shaped caudal fins are diagnostic. The male spine 4.5 (7.6) in head; second anal spine 3.9 (8.7) in of C. johnsoni is primarily orange-red with bright red head; third anal spine 3.4 (10.1) in head; fourth and fifth median fins. The new species appears to be most anal soft rays longest, 2.4 (14.2) in head; caudal fin closely related to C. lunatus (Figure 2). The two species lunate with protruding upper and lower lobes, its length have a similar dorsal fin shape and there are remark - 1.1 (30.3) in head; caudal concavity 2.4 (14.2) in head; able similarities in the coloration of the median fins, third and fourth pectoral rays longest, 1.5 (22.9) in head; which are dark (brown in C. brunneus and black in C. origin of pelvic fins below pectoral in base; pelvic fin lunatus ) with a broad white margin posteriorly on the length 1.5 (22.7) in head, the pelvic fin tips falling well dorsal and caudal fins. However, C. lunatus differs in short of anal fin origin when depressed; length of pelvic having a broad orange-yellow zone along the sides at fin spine 3.0 (11.2) in head. the level of the pectoral fins and fish from the Ryukyu Colour of male holotype when fresh (Fig. 1): mainly Islands have a purplish coloration on the sides, which dark brown with slight purplish hue; breast and belly angles upward posteriorly to encompass the entire cau - with bronze hue, a diamond shaped, red-brown marking dal peduncle. Male specimens of C. lunulatus from the on each scale; iris bright red and pupil blackish; dorsal Ogasawara Islands have pinkish coloration on the lower fin dark brown with white distal margin that gradually sides. Males from both the Ryukyu and Ogasawara widens posteriorly, resulting in a broad white area islands differ from those of C. brunneus in having a pale around angular posteriormost portion of fin; anal fin dark yellowish band along the base of the dorsal fin and brown with narrow white margin; caudal fin mainly irregular diagonal red lines on the snout and postorbital brown including prominent upper and lower margins of portion of the head that extend onto the anterodorsal fin, the posterior margin broadly white; pelvic fins dark part of the body. brown; pectoral rays translucent with dark brown base. There is also a possible difference in the number of gill

Fig. 2. Cirrhilabrus lunatus, freshly collected male holo - Fig. 3 . Cirrhilabrus lunatus, freshly collected female type, 67.7 mm SL, Okinawa, Ryukyu Islands. Photo by J. paratype, 45.1 mm SL, Ani-jima, Ogaswara Islands. Photo E. Randall. by J. E. Randall.

3 aqua vol. 11 no. 1 - 2006 Cirrhilabrus brunneus , a new wrasse (Pisces: Labridae) from north-eastern Kalimantan, Indonesia rakers on the first branchial arch. Randall and Masuda from Fiji. aqua, Journal of Ichthyology and Aquatic (1991) indicated a range of 14-17 gill rakers for six spec - Biology, 7 (3): 103-112. imens of C. lunatus in comparison to 13 rakers for the Randall, J. E. 1988. Five new wrasses of the genera holotype of C. brunneus . Of course, it would be desir - Cirrhilabrus and Paracheilinus (Perciformes: Labridae) able to count additional specimens of the latter species from the Marshall Islands. Micronesica, 21: 199-226. to verify this disparity. There is also a substantial differ - Randall, J. E. & H. Masuda, 1991. Two new labrid ence in maximum size. Males of C. lunatus attain a fishes of the genus Cirrhilabrus from Japan. Revue maximum standard length of at least 84.9 mm. In com - française d’Aquariologie, 18 (2): 53-60. parison, the holotype of C. brunneus, measuring 43.6 Randall, J.E. & B.H. Nagareda. 2002. Cirrhilabrus mm SL, is scarcely more than half this size. Moreover, bathyphilus , a new deep-dwelling labrid fish from the four more males of similar size were sighted at the type Coral Sea. Cybium, 26 (2): 123-127. locality. Randall, J.E. & R.L. Pyle. 2001. Three new species of Several initial phase (female) individuals of C. brun - labrid fishes of the genus Cirrhilabrus from islands of neus were observed at the type locality. They were very the tropical Pacific. aqua, Journal of Ichthyology and similar in appearance to the female of C. lunatus from Aquatic Biology, 4 (3): 89-98. the Ogasawara Islands that was illustrated by Randall Senou, H. and T. Hirata. 2000. A new labrid fish, Cir - and Masuda (1991) and is included here as Fig. 3. rhilabrus katoi , from southern Japan. Ichthyological Cirrhilabrus brunneus is presently known only from the Research, 4 (1): 89-93. type locality at north-eastern Kalimantan. However, it is likely to occur in adjacent areas, particularly the Palawan region of the Philippines, which frequently shares faunal elements with northern Borneo. The species was sighted on only one occasion despite nearly one month of diving. However, very few dives involved a habitat situation similar to that of the type locality, which was characterised by a steep slope beginning in relatively deep water. About 10 individuals, including the eventual holotype, were observed in 40 m depth on a flat to gently sloping rubble bottom along the upper edge of a vertical dropoff.

Etymology This species is named brunneus (Latin: dusky or dark) with reference to the overall colour pattern of the holo - type.

Acknowledgements I am very grateful to The Nature Conservancy (TNC), especially Scott Stanley and Budy Wiryawan, for inviting me to participate in the Borneo Rapid Eco - logical Assessment. Excellent diving assistance was provided by TNC staff members Ibu Katherina, Tasrif Kartawijaya, Handoko Adi Susanto, and Irfan Yulianto. I am also grateful to Pak Witir, skipper of the TNC vessel “Ridges to Reefs” and his able assistant Tony Suhaimi. Coral specialist Emre Turak provided companionship and valuable advice throughout the survey. I also thank Suzette Stephens of TNC for logistic coordination. The manuscript was kindly reviewed by John E. Randall.

References Allen, G. R. 2000. Description of a new wrasse (Pisces: Labridae: Cirrhilabrus) from northern Sumatra, Indonesia. aqua, Journal of Ichthyology and Aquatic Biology, 4 (2): 45-50. Allen, G. R., J. E. Randall & B. A. Carlson. 2003. Cirrhilabrus marjorie , a new wrasse (Pisces: Labridae) aqua vol. 11 no. 1 - 2006 4 aqua, Journal of Ichthyology and Aquatic Biology

Redescription of Kryptolebias ocellatus (Hensel) and K. caudomarginatus (Seegers) (Teleostei: Cyprinodontiformes: Rivulidae), two killifishes from mangroves of south-eastern Brazil

Wilson J. E. M. Costa

Laboratório de Ictiologia Geral e Aplicada, Departamento de Zoologia, Universidade Federal do Rio de Janeiro, Caixa Postal 68049, CEP 21944-970, Rio de Janeiro, Brasil. E-mail: [email protected]

Accepted: 17.01.06

Keywords de colorido exclusivos em macho. A localidade tipo de Fish, Cyprinodontiformes, Rivulidae, Kryptolebias , K. caudomarginatus é corrigida. Neotropica, mangrove, systematics, hermaphrodite, morphology Zusammenfassung Die systematische Stellung von Kryptolebias ocellatus Abstract (Hensel) und K. caudomarginatus (Seegers) wurde Taxonomy of Kryptolebias ocellatus (Hensel) and K. bisher nicht ausreichend begründet und ihre Beschrei - caudomarginatus (Seegers) has been poorly defined bung beruhte auf einigen wenigen Exemplaren aus and mostly based on a few specimens bred in aquaria. Aquarien. Sie werden hier zu gültigen Arten erklärt und They are considered valid species and are redescribed auf der Grundlage neuerer Fänge im Gebiet der Typus - based on recent collections in the type locality area, the lokalität neu beschrieben, den Mangrovensümpfen der mangroves of Rio de Janeiro state, south-eastern Region Rio de Janeiro, in SO-Brasilien. Die beiden Arten Brazil. Both species are considered closely related to K. werden als nahe verwandt mit K. marmoratus (Poey) marmoratus (Poey), with which they share the presence eingestuft; gemeinsame Merkmale sind vier Neuromas - of four neuromasts on the posterior supraorbital series, ten in der hinteren supraorbitalen Reihe, ein langes a long anterior nostril, and a bony laminar ventral vorderes Nasenloch und ein knochiger, blättchenhafter process on the fifth ceratobranchial; K. ocellatus is dis - ventraler Fortsatz am fünften Ceratobranchiale; K. ocel - tinguished by a unique colour pattern in the hermaphro - latus unterscheidet sich durch ein unverkennbares Farb - dite, more slender caudal peduncle, and a shorter pelvic muster ohne Geschlechtsunterschied, einen schlan- fin, and K. caudomarginatus by possessing more keren Schwanzstiel und eine kürzere Bauchflosse, vomerine teeth, longer dorsal fin base, and a unique während K. caudomarginatus mehr Schlundzähne, eine male colour pattern. The type locality of K. caudomar - längere Rückenflossenbasis und ein unverkennbares ginatus is coorrected. männliches Farbmuster besitzt. Die Typuslokalität von K. caudomarginatus wird korrekt abgegrenzt. Resumo A taxonomia de Kryptolebias ocellatus (Hensel) e K. Résumé caudomarginatus (Seegers) tem sido fracamente La taxinomie des espèces Kryptolebias ocellatus definida, principalmente com base em poucos (Hensel) et K. caudomarginatus (Seegers) a été peu espécimes criados em aquários. Elas são consideradas spécifiée et basée surtout sur un petit nombre de spéci - espécies válidas e são redescritas com base em mens élevés en aquarium. Elles sont considérées coleções recentes na área da localidade tipo, os comme des espèces valides et sont redécrites sur base manguezais do estado do Rio de Janeiro, sudeste do de collectes récentes dans la région de la localité-type, Brasil. Ambas espécies são consideradas estreita - les mangroves de l'état de Rio de Janeiro, au sud-est du mente aparentadas a K. marmoratus (Poey), com a Brésil. Les deux espèces paraissent très proches de K. qual elas compartilham a presença de quatro neuro - marmoratus (Poey), et, comme celle-ci, montrent la mastos na série supraorbital posterior, narina anterior présence de quatre neuromastes sur la série longa, e processo ventral laminar ósseo em quinto cer - postérieure supra-orbitale, une longue narine antérieure atobranquial; K. ocellatus é distinguível por possuir um et une excroissance osseuse laminaire ventrale sur la padrão de colorido exclusivo em hermafrodita, pedún - cinquième cératobranchiale. K. ocellatus se distingue culo caudal mais esguio e nadadeira pélvica mais curta, par une coloration unique du pédoncule caudal her - e K. caudomarginatus por possuir mais dentes no maphrodite, plus mince et une pelvienne plus courte, et vomer, base de nadadeira dorsal mais longa e padrões K. caudomarginatus, par un plus grand nombre de dents

5 aqua vol. 11 no. 1 - 2006 Redescription of Kryptolebias ocellatus (Hensel) and K. caudomarginatus (Seegers) (Teleostei: Cyprinodontiformes: Rivulidae) vomériennes, une plus longue base de la dorsale et un meulen & Hrbek, 2005) support monophyly of the latter patron de coloration unique pou le mâle. La localité-type assemblage. Among species of this assemblage, K. de K. caudomarginatus est correctement identifiée. ocellatus and K. marmoratus are self-fertilizing her - maphroditic species, a condition unique among verte - Sommario brates ( e. g. , Harrington, 1961). Hermaphrodites are La tassonomia di Kryptolebias ocellatus (Hensel) e female-like individuals, exhibiting typical female colour K. caudomarginatus (Seegers) è poco definita e patterns. Populations are mainly composed of her - basata per lo più su pochi esemplari provenienti da maphrodites, while functional females are unknown, and accoppiamenti in acquario. Esse sono considerate males are uncommon or very rare ( e. g. , Davis et al. , specie valide e vengono qui descritte nuovamente 1990; Turner et al. , 1992). For example, Taylor (1990) sulla base di recenti catture nella località tipo, le man - collected over 250 specimens of K. marmoratus in grovie dello stato di Rio de Janeiro, Brasile sud-orien - Florida, but none of them was a male. tale. Entrambe sono considerate molto vicine a Although some species of Kryptolebias are among the K. marmoratus (Poey), con cui condividono la pre - oldest described species of the Rivulidae, their taxon - senza di quattro neuromasti sulla serie supraorbitale omy is still confused. The best example is K. ocellatus , posteriore, una lunga narice anteriore e un processo described by Hensel (1868) based on a single specimen laminare ventrale osseo sul quinto ceratobrachiale; collected in Rio de Janeiro. The identity of K. ocellatus K. ocellatus si distingue per una tipica colorazione was difficult to determine between 1906 and 1984, a nell’ermafrodita, un peduncolo caudale più sottile e le period when K. caudomarginatus , another species from pinne pelviche più corte, mentre K. caudomarginatus Rio de Janeiro and undescribed at that time, was usu - si differenzia per possedere più denti vomerini, una ally identified as Rivulus ocellatus (= K. ocellatus ). pinna dorsale a base più lunga e una singolare colo - Seegers (1984) established the identity of K. ocellatus razione nel maschio. La località tipo K. caudomargi - and first described K. caudomarginatus based upon natus è definita in modo corretto. aquarium bred specimens. Seegers also considered K. marmoratus conspecific to K. ocellatus , proposing the Introduction two names as subspecies of K. ocellatus , which would Kryptolebias Costa was recently established to include have chronological priority over K. marmoratus , some species previously placed in Rivulus : K. brasilien - described 12 years later (Poey, 1980). However, K. mar - sis (Valenciennes), K. caudomarginatus (Seegers), and moratus is an important experimental fish for researches K. ocellatus (Hensel) from south-eastern Brazil; K. on carcinogenicity, mutagenesis, teratogenesis, and campelloi (Costa) from eastern Brazilian Amazon; K. other scientific areas, in which the specific epithet mar - marmoratus (Poey), recorded from a broad geographic moratus is consistently applied, whereas the specific range, including the south-eastern United States, epithet ocellatus , the putative senior synonym of mar - Bahamas, Yucatan Peninsula, Cuba and other moratus , was poorly known and never correctly applied Caribbean islands, Venezuela and Guianas (Costa, for many decades. These were the basic arguments 2004a, 2004b). Three poorly known nominal species, used by Lazara & Smith (1990) to propose suppression considered synonyms of K. marmoratus (e. g. , Costa, of the specific epithet ocellatus and conservation of the 2003), were also included in Kryptolebias : K. bonairen - name marmoratus . However, Costa (1994) did not sis (Hoedeman) from Bonaire, Curação, and coastal agree with such a proposal, which would be premature, Venezuela; K. heyei (Nichols) from Isla Saona, Domini - since no rigorous taxonomic study was developed to can Republic; and, K. garciai (De la Cruz & Dubitsky) check whether ocellatus and marmoratus were in fact from Cuba (Costa, 2004a). Subsequently, Vermeulen & the same species. The International Commission of Hrbek (2005) described K. sepia from Surinam. Zoological Nomenclature (1995) then decided to give Kryptolebias has been hypothesized to be the sister precedence to the name marmoratus over the name group to a clade comprising all other rivulids on the basis ocellatus , whenever the two names are considered to be of morphological and some molecular analyses (Hrbek synonyms. However, due to difficulties in collecting & Larson, 1999; Costa, 2004a, 2004c; Vermeulen & small fishes in a mangrove habitat, K. ocellatus and K. Hrbek, 2005) or a basal clade within a monophyletic lin - caudomarginatus have been insufficiently sampled in eage including all rivulids except cynolebiatines accord - scientific collections, and consequently are still poorly ing to other molecular analyses (Murphy et al. , 1999; known taxonomically. Both species are herein Vermeulen & Hrbek, 2005). Kryptolebias brasiliensis , K. redescribed, based on recent collections made in their campelloi , and K. sepia are similar to most rivulids in type locality areas. inhabiting freshwater pools and streams ( e. g. , Costa, 2004a; Vermeulen & Hrbek, 2005), while K. caudomar - Material and methods ginatus , K. ocellatus , and K. marmoratus live in brackish All material was collected with dip nets. Measurements to salt water, usually in mangrove swamps ( e. g. , Taylor, and counts follow Costa (1995). Measurements are pre - 1988; the present paper). Both morphological (Costa, sented as percentages of standard length (SL), except 2004a) and molecular studies (Murphy et al. , 1999; Ver - those related to head morphology, and are expressed aqua vol. 11 no. 1 - 2006 6 Wilson J. E. M. Costa as percentages of head length. Fin ray counts include all and K. caudomarginatus in having long anterior nostril, elements. Numbers of vertebrae, gill rakers, and pec - its tip extending beyond anterior profile of the head. Also toral, pelvic and caudal fin rays were recorded only from similar to K. marmoratus and K. caudomarginatus by cleared and stained specimens. The compound caudal possessing numerous scales in the longitudinal series centrum was counted as a single element. Osteological (44-48, vs. 30-32 K. brasiliensis and K. campelloi , and preparations were made according to Taylor and Van 38-40 in K. sepia ) and four neuromasts on the posterior Dyke (1985). Terminology for frontal squamation and supraorbital series ( vs. three). It differs from K. mar - the cephalic neuromast series follow Hoedeman (1958) moratus and K. caudomarginatus in having a vertical and Costa (2001), respectively. The abbreviation “c&s” row of pale silver to pale gold dots on the dorsal portion stands for specimens cleared and stained for bone and of flank in the hermaphrodite ( vs. pale silver spots over cartilage. Material is deposited in Universidade Federal the whole flank in the hermaphrodite of K. marmoratus do Rio de Janeiro, Rio de Janeiro, Brazil (UFRJ). Other and in the female of K. caudomarginatus ), more slender abbreviations for of institution names are: ZFMK, Zoolo - caudal peduncle (caudal peduncle depth 12.3-13.4 % gisches Farschungsinstitut und Museum Alexander SL, vs. 13.7-16.3 % SL), and shorter pelvic fin (pelvic fin Koenig, Bonn, and ZMB, Zoologisches Museum, Hum - length 6.0-7.8 % SL, vs. 8.5-10.7 % SL). boldt-Universität, Berlin. Description Kryptolebias ocellatus (Hensel) The description is based on hermaphrodites only. Mor - (Figs. 1-2; Table I) phometric data appear in Table I. Largest specimen 49.8 mm SL. Dorsal profile approximately straight to Rivulus ocellatus Hensel, 1868: 365 (type locality: Rio slightly convex from snout to end of dorsal fin base, de Janeiro; holotype: ZMB 7448). approximately straight on caudal peduncle. Ventral pro - file gently convex from lower jaw to end of anal fin base, Material examined about straight on caudal peduncle. Body slender, sub - Brazil: Estado do Rio de Janeiro: UFRJ 1806, 3; man - cylindrical anteriorly, about as deep as wide, to com - grove swamp near Itaguaí; M. S. Melgaço, 25 Jul. 1991. pressed posteriorly. Greatest body depth at vertical just UFRJ 6234, 4 (c&s); mangrove canal, near EMBRAPA, in front to pelvic fin base. Jaws short, snout blunt. Ante - rio Piracão basin, a tributary to baía de Sepetiba, rior naris cylindrical and long, extending beyond anterior Guaratiba; W. J. E. M. Costa, C. P. Bove & B.B. Costa, profile of the head. Urogenital papilla pocket-shaped. 2004. UFRJ 6237, 3; idem ; W. J. E. M. Costa, C. P. Tip of dorsal and anal fins rounded. Anal fin rays 4 and Bove & T. Litz, 19 Feb. 2005. UFRJ 6243, 3; idem ; W. 5 longer than other anal fin rays. Caudal fin rounded. J. E. M. Costa, J. Barata, A. Moreira & J. P. Moreira, 14 Pectoral fin short and rounded, posterior margin reach - Aug. 2005. UFRJ 6252, 9; idem ; W. J. E. M. Costa, J.L. ing vertical just posterior to midlength between pectoral Mattos, L. Villa Verde, F. Ottoni & E. Mattos, 22 Oct. fin and pelvic fin bases. Pelvic fin elliptical, tip reaching 2005. Material collected in mangrove swamp near Cax - base of 1st anal fin ray. Pelvic fin bases medially sepa - ias, baía de Guanabara, was not preserved. rated by short interspace. Dorsal fin origin on vertical between base of 10th and 11th anal fin rays, and Diagnosis between neural spines of 18th and 20th vertebrae. Anal Distinguished from all rivulids except K. marmoratus fin origin between pleural ribs of 14th and 15th verte -

Fig. 1. Kryptolebias ocellatus , hermaphrodite, UFRJ 6243, 46.8 mm SL; Brazil: Rio de Janeiro: Guaratiba. Photo by W. J. E. M. Costa.

7 aqua vol. 11 no. 1 - 2006 Redescription of Kryptolebias ocellatus (Hensel) and K. caudomarginatus (Seegers) (Teleostei: Cyprinodontiformes: Rivulidae)

Table I. Morphometric data of Kryptolebias ocellatus and K. caudomarginatus .

K. ocellatus K. caudomarginatus hermaphrodites males females (n = 10) (n = 10) (n = 10) Standard length (mm) 27.2-48.3 28.3-48.5 27.7-36.3 Percent of standard length Body depth 17.5-19.7 19.2-23.4 19.4-23.3 Caudal peduncle depth 12.3-13.4 13.7-16.3 13.9-16.1 Predorsal length 73.5-77.2 72.5-76.3 72.1-78.0 Prepelvic length 56.9-59.9 56.9-59.4 57.6-61.5 Length of dorsal fin base 9.2-10.7 11.1-13.5 10.9-12.5 Length of anal fin base 14.1-15.6 15.1-17.4 14.4-16.5 Caudal fin length 27.9-31.4 29.2-34.4 30.2-32.8 Pectoral fin length 17.4-19.0 17.1-20.4 18.4-20.8 Pelvic fin length 6.0-7.8 8.7-9.5 8.5-10.7 Head length 24.0-26.0 24.8-27.7 24.8-27.8 Percent of head length Head depth 62.8-72.1 60.3-78.2 64.0-77.8 Head width 75.0-93.6 74.6-91.6 76.5-89.7 Snout length 13.5-15.7 13.5-15.5 14.0-15.9 Lower jaw length 20.2-25.0 18.4-25.7 19.1-24.1 Eye diameter 28.4-33.0 31.2-35.9 32.2-36.0

brae. Dorsal fin rays 8-9; anal fin rays 11-12; caudal fin rays 30-31; pectoral fin rays 13; pelvic fin rays 6. Scales small, cycloid. Body and head entirely scaled, except anterior ventral surface of head. Body squama - tion extending over anterior 40 % of caudal fin base; no scales on dorsal and anal fin bases. Frontal squamation E-patterned; E-scales overlapping medially; scales arranged in regular circular pattern around A-scale with - out exposed margins. Longitudinal series of scales 47- 48; transverse series of scales 13-14; scale rows around caudal peduncle 24-26. Contact organs absent. Cephalic neuromasts: supraorbital 3 + 4, parietal 1-2, anterior rostral 1, posterior rostral 1, infraorbital 1 + 1 + 13-14 + 1, preorbital 3, otic 1, post-otic 2, supratempo - ral 1, median opercular 1, ventral opercular 2, preoper - cular 2 + 7-8, mandibular 4 + 2-3, lateral mandibular 4. Interhyal ossified. Rostral cartilage longer than wide, width about 60% length. Basihyal subtriangular, width about 65 % length; basihyal cartilage about 10 % of basihyal length. Six branchiostegal rays. Second pharyngobranchial teeth 4-5. Gill rakers of first branchial arch 1 + 8. Vomerine teeth 1-4. Ventral process of post - temporal long. Total vertebrae 33-34. Coloration in life: Side of body light brown with row of pale silver to pale golden dots on dorsal portion of flank; black humeral blotch with narrow anterior extension; round black spot with bright grey margin on dorsal end of caudal peduncle, sometimes other black spot below. Fig. 2. Kryptolebias ocellatus , hermaphrodite, UFRJ Dorsum light brown. Venter white. Side of head light 6237, 36.8 mm SL; Brazil: Rio de Janeiro: Guaratiba. brown on dorsal portion, pale silver with great concen - Photo by W. J. E. M. Costa. tration on melanophores ventrally. Lower jaw grey. Iris aqua vol. 11 no. 1 - 2006 8 Wilson J. E. M. Costa brown. Unpaired fins grey with hyaline small spots on from Rio de Janeiro, identified as “primary male of ocel - basal portion; anal fin with pale yellow subdistal stripe, latus”. Notare’s colour photo shows a fish in which the and distal grey stripe. Pelvic and pectoral fins hyaline. flank is light grey with dark red spots. There is a dark grey humeral spot and a grey caudal peduncle spot, Distribution and habitat and, the unpaired fins have a pale yellow subdistal zone Kryptolebias ocellatus inhabits the shallow parts and a dark grey to black distal zone. This material was (about 10-40 cm deep) of mangrove swamps associ - not preserved for study, and its identification needs con - ated with Sepetiba and Guanabara bays (Fig. 3). It is firmation. always sympatric to K. caudomarginatus , which is con - spicuously more abundant, and sometimes with two Kryptolebias caudomarginatus (Seegers) species of poeciliids, Poecilia vivipara Bloch & Schnei - (Figs. 4-7; Table I) der and Phalloptychus januarius (Hensel), an eleotri - dine gobiid, Dormitator maculatus (Bloch), and the cal - Rivulus caudomarginatus Seegers, 1984: 307 (type lichthyid Callichthys callichthys (Linnaeus). It is never locality: Greta Funda [correctly Grota Funda, an equiv - found in freshwater streams. The occurrence of this ocal type locality as discussed in “Remarks” below] species has already been reported in Santos, São near a technical centre of the Army, southern Rio de Paulo (Huber, 1992), but the material from this region Janeiro [correctly near Technological Center of the was not examined in the present paper. Brazilian Army, Guaratiba, south-western Município do Rio de Janeiro, Estado do Rio de Janeiro, south - Remarks eastern Brazil, rio Piracão basin, a tributary to Baía de All the material herein examined included only her - Sepetiba, 23°00’18.1”S 43°33’34.9”W, altitude 8 m]; maphrodites. Three recent collecting trips were unsuc - holotype: ZFMK 12848). cessful in collecting males. However, Notare (1988) pro - vided a colour photo of a live male identified as “Rivulus Material examined caudomarginatus var. (?)” collected in Rio de Janeiro Brazil: Estado do Rio de Janeiro: UFRJ 1805, 32; man - and Huber (1992) illustrated a similar live male, also grove near Itaguaí; M.S. Melgaço, 25 Jul. 1991. UFRJ 1047, 22; UFRJ 3583, 3 (c&s); Madeira, mangrove swamp or mangrove habitat near Itaguaí; M. S. Mel - gaço, 25 Jul. 1991. UFRJ 4316, 30; rio Iriri, a tributary to Baía de Guanabara, Barão de Iriri; W. J. E. M. Costa, F. Pupo, E. Araujo & F. Autran, 4 Dec. 1998. UFRJ 6235, 15; UFRJ 6236, 6 (c&s); mangrove canal, near EMBRAPA, rio Piracão basin, a tributary to Baía de Sepetiba, Guaratiba; W. J. E. M. Costa, C. P. Bove & B.B. Costa, 2004. UFRJ 6238, 4; idem ; W. J. E. M. Costa, C. P. Bove & T. Litz, 19 Feb. 2005. UFRJ 6244, 12; idem ; W. J. E. M. Costa, J. Barata, A. Moreira & J. P. Moreira, 14 Aug. 2005. UFRJ 6253, 4; idem ; W. J. E. M. Costa, J. L. Mattos, L. Villa Verde, F. Ottoni & E. Mat - tos, 22 Oct. 2005. Material collected in mangrove habi - tat near Caxias, Baía de Guanabara, was not preserved.

Diagnosis Distinguished from all rivulids except K. marmoratus and K. ocellatus in having long anterior nostril, its tip extending beyond anterior profile of the head. Also sim - ilar to K. marmoratus and K. ocellatus by possessing numerous scales in the longitudinal series (44-48, vs. 30-32 K. brasiliensis and K. campelloi, and 38-40 in K. sepia in other species of Kryptolebias) and four neuro - masts on the posterior supraorbital series ( vs. three). It differs from K. marmoratus and K. ocellatus in having more vomerine teeth (6-17, vs. 1-4), longer dorsal fin base (10.9-13.5 % SL, vs. 9.2-10.7 % SL), and a unique colour pattern of flank and caudal fin in male (flank with Fig. 3. Mangrove at Guaratiba, Rio de Janeiro, Brazil, black oblique bars on its posterior half, vs. dark red habitat of Kryptolebias ocellatus and K. caudomargina - spots; caudal fin with a broad white to yellowish white tus . Photo by W. J. E. M. Costa. subdistal zone, strongly contrasting with a broad black

9 aqua vol. 11 no. 1 - 2006 Redescription of Kryptolebias ocellatus (Hensel) and K. caudomarginatus (Seegers) (Teleostei: Cyprinodontiformes: Rivulidae)

of 8th and 10th anal fin rays, and between neural spines of 17th and 19th vertebrae. Anal fin origin between pleural ribs of 13th and 15th vertebrae. Dorsal fin rays 8-10; anal fin rays 11-13; caudal fin rays 29-32; pec - toral-fin rays 13; pelvic fin rays 5-6. Scales small, cycloid. Body and head entirely scaled, except anterior ventral surface of head. Body squama - tion extending over anterior 40 % of caudal fin base; no scales on dorsal and anal fin bases. Frontal squamation E-patterned; E-scales overlapping medially; scales arranged in regular circular pattern around A-scale with - out exposed margins. Longitudinal series of scales 43- 48; transverse series of scales 9-10; scale rows around caudal peduncle 20-22. Contact organs absent. Fig. 4. Kryptolebias caudomarginatus , male, UFRJ Cephalic neuromasts: supraorbital 3-4 + 4, parietal 2, 6235, 34.8 mm SL; Brazil: Rio de Janeiro: Guaratiba. anterior rostral 1, posterior rostral 1, infraorbital 1 + 2 + Photo by W. J. E. M. Costa. 12-13 + 1, preorbital 3, otic 1, postotic 2, supratemporal 1, median opercular 1, ventral opercular 2, preopercular 2 + 6-7, mandibular 5 + 2, lateral mandibular 3-5. Interhyal ossified. Rostral cartilage longer than wide, width about 60 % length. Basihyal subtriangular, width about 50 % length; basihyal cartilage about 10 % basi - hyal length. Six branchiostegal rays. Second pharyngo - branchial teeth 3-5. Gill rakers of first branchial arch 1-2 + 8-9. Vomerine teeth 6-17. Ventral process of post - temporal long. Total vertebrae 32-33.

Fig. 5. Kryptolebias caudomarginatus , male, UFRJ 6238, 42.4 mm SL; Brazil: Rio de Janeiro: Guaratiba. Photo by W. J. E. M. Costa. distal zone, vs. indistinct pale yellow subdistal zone bor - dered by narrow black distal zone).

Description Morphometric data appear in Table I. Male larger than female, largest male 54.6 mm SL. Dorsal profile slightly convex from snout to end of dorsal fin base, approxi - mately straight on caudal peduncle. Ventral profile gen - tly convex from lower jaw to end of anal fin base, about straight on caudal peduncle. Body slender, subcylindri - Fig. 6. Kryptolebias caudomarginatus , female, UFRJ cal anteriorly, about as deep as wide, to compressed 6235, 31.9 mm SL; Brazil: Rio de Janeiro: Guaratiba. posteriorly. Greatest body depth at vertical just in front Photo by W. J. E. M. Costa. of pelvic fin base. Jaws short, snout blunt. Anterior naris cylindrical and long, extending anteriorly beyond ante - rior profile of the head. Urogenital papilla globular in male, pocket-shaped in female. Tip of dorsal and anal fins rounded. Anal fin rays 4 and 5 longer than other anal fin rays. Caudal fin rounded. Pectoral fin short and rounded, posterior margin reach - ing vertical just posterior to midlength between pectoral fin and pelvic fin bases. Pelvic fin elliptical, tip reaching between base of 2nd and 4th anal fin rays in male, and between base of 1st and 2nd anal-fin rays in female. Fig. 7. Kryptolebias caudomarginatus , female, UFRJ Pelvic fin bases medially separated by short interspace 6238, 36.3 mm SL; Brazil: Rio de Janeiro: Guaratiba. or in contact. Dorsal fin origin on vertical between base Photo by W. J. E. M. Costa. aqua vol. 11 no. 1 - 2006 10 Wilson J. E. M. Costa

Coloration in life: Male: Side of body pale bluish sil - and K. caudomarginatus were made in a mangrove ver; black spots irregularly shaped and arranged on area near the Technological Center of the Brazilian anterior half of flank and black oblique bars, usually Army, Guaratiba, south-western part of the metropol - zigzag shaped, sometimes broken, forming isolated itan region of the Rio de Janeiro city, by aquarists. black spots on posterior half of flank; approximately rec - Seegers (1984) mentioned that the type specimens of tangular black humeral blotch; round black spot on dor - R. caudomarginatus were the progeny of fish col - sal end of caudal peduncle, sometimes other black spot lected in “Greta Funda”, a misspelling of Grota Funda, below. Dorsum light brown, with green iridescence of near a technical center of the Army. However, Grota laterodorsal scales. Venter white. Side of head light Funda is situated about 4 km east of the Technologi - greenish brown on dorsal portion, pale silver with great cal Center of the Brazilian Army, at a small coastal hill concentration of melanophores ventrally. Lower jaw (Serra Geral de Guaratiba), at an altitude of 200-300 grey. Iris brown. Dorsal and anal fins light grey with m. No habitat suitable to the occurrence of rivulids is small dark grey to black small spots, broad subdistal present in Grota Funda, thus it is considered an white stripe and narrow distal black stripe. Caudal fin equivocal type locality. dark grey, with broad white to yellowish-white subdistal zone and broad black distal zone; often dorsal and ven - Discussion tral margins and filaments dark orange. Pelvic fin hya - Monophyly of the assemblage including K. ocellatus , line with subdistal white line and black margin. Pectoral K. marmoratus and K. caudomarginatus has been sup - fin hyaline. ported by different studies (e. g. , Murphy et al. , 1999; Female: Side of body light brown with pale silver spots; Costa, 2004a; Vermeulen & Hrbek, 2005). New evi - black humeral blotch; round black spot with bright grey dence corroborating this hypothesis is herein provided: margin on dorsal end of caudal peduncle, Caudal fin 1) presence of four neuromasts on the posterior hyaline, with small faint grey spots on basal portion and supraorbital series (Fig. 8), a condition not occurring pale grey margin. Pelvic and pectoral fins hyaline. in other species of Kryptolebias nor in members of All described color patterns visible in preserved speci - other basal rivulid lineages, such as the genera Pror - mens, both in male and female. ivulus Costa and Rivulus Poey, which have three neu - romasts; Distribution and habitat 2) long anterior nostril (Fig. 8), a condition not found As described for K. ocellatus . in other aplocheiloid fishes; 3) bony laminar ventral process close to the lateral Remarks edge of the fifth ceratobranchial (Fig. 9), not found During the 1980’s, several collections of K. ocellatus elsewhere among aplocheiloid fishes.

Fig. 8. Diagrammatic representation of the head, dorsal Fig. 9. Fifth ceratobranchial, ventral view, of Kryptolebias view, of Kryptolebias caudomarginatus , UFRJ 6253, caudomarginatus , UFRJ 6236, male, 38.5 mm SL. male, 48.5 mm SL. Drawn by the author Drawn by the author

11 aqua vol. 11 no. 1 - 2006 Redescription of Kryptolebias ocellatus (Hensel) and K. caudomarginatus (Seegers) (Teleostei: Cyprinodontiformes: Rivulidae)

Acknowledgements Hensel, R. 1868. Beiträge zur Kenntnis der Wirbelthiere I am grateful to C. P. Bove, B. B. Costa, T. Litz, J. P. Südbrasiliens. Archiv für Naturgeschichte Wiegman , Moreira, A. Moreira, J. Barata, J. L. Mattos, L. Villa 34 : 323-375. Verde, F. Ottoni, and E. Mattos for help during collec - Hoedeman, J. J. 1958. The frontal scalation pattern in tions. This study was also supported by CNPq (Con - some groups of toothcarps (Pisces, Cyprinodontif - selho Nacional de Desenvolvimento Científico e Tec - ormes). Bulletin of Aquatic Biology , 1: 23-28. nológico – Ministério de Ciência e Tecnologia) and Hrbek, T. & A. Larson. 1999. The diapause in the killi - FAPERJ (Fundação de Amparo à Pesquisa do Estado fish family Rivulidae (Atherinomorpha, Cyprinodontif - do Rio de Janeiro). ormes): a molecular phylogenetic and biogeographic perspective. Evolution , 53 : 1200-1216. Huber, J. H. 1992. Review of Rivulus: ecobiogeografia References - relationships. Cybium, Société Française d’Ichtyolo - gie, Paris, 572 + 13 pp. Costa, W. J. E. M. 1994. Comments on the proposed International Commission of Zoological Nomencla - conservation of the specific name of Rivulus mar - ture. 1995 Rivulus marmoratus Poey, 1880 (Oste - moratus Poey, 1880 (Osteichthyes, Cyprinodontif - ichthyes, Cyprinodontiformes): given precedence over ormes). Bulletin of Zoological Nomenclature , 51 : 46- R. ocellatus Hensel, 1868, and a neotype designated 47. for R. marmoratus . Bulletin of Zoological Nomencla - Costa, W. J. E. M. 1995. Pearl killifishes - the Cynolebi - ture , 52 : 106-108. atinae: systematics and biogeography of the neotropi - Lazara, K. J. & M. L. Smith 1990 Rivulus marmoratus cal annual fish subfamily. TFH, Neptune City, 128 pp. Poey, 1880 (Osteichthyes, Cyprinodontiformes): pro - Costa, W. J. E. M. 1998. Phylogeny and classification posed conservation of the specific name. Bulletin of of Rivulidae revisited: evolution of annualism and Zoological Nomenclature , 47 : 191-194. miniaturization in rivulid fishes (Cyprinodontiformes: Murphy, W. J., J. E. Thomerson & G. E. Collier. 1999. Aplocheiloidei). Journal of Comparative Biology , 3: 33- Phylogeny of the neotropical killifish family Rivulidae 92. (Cyprinodontiformes, Aplocheiloidei) inferred from Costa, W. J. E. M. 2001 . The neotropical annual fish mitochondrial DNA sequences. Molecular and Phylo - genus Cynolebias (Cyprinodontiformes: Rivulidae): genetic Evolution , 13 : 289-301. phylogenetic relationships, taxonomic revision and Notare, M. 1988. Peixes raros do Brasil. Revista de biogeography. Ichthyological Exploration of Freshwa - Aquariofilia , 5: 52-53. ters , 12 : 333-383. Poey, F. 1880. Revisio piscium cubensium. Anales de Costa, W. J. E. M. 2003 . Family Rivulidae (South Amer - la Sociedad Española de Historia Natural , 9: 243-261. ican annual fishes). In : R.E. Reis, Kullander, S.O. & Seegers, L. 1984 Zur Revision der Rivulus -Arten Ferraris, C.J., Jr. (eds) Check list of the freshwater Südost-Brasiliens, mit einer Neubeschreibung von fishes of South and Central America , Edipucrs, Porto Rivulus luelingi n. sp. und Rivulus caudomarginatus n. Alegre, pp 526-548. sp. (Pisces: Cyprinodontidae: Rivulinae). Zoologische Costa, W. J. E. M. 2004a. Relationships and redescrip - Beiträge , 28 : 271-320. tion of Fundulus brasiliensis (Cyprinodontiformes: Taylor, S. 1988. Observations on the ecology of the kil - Rivulidae), with description of a new genus and notes lifish Rivulus marmoratus (Cyprinodontidae) in an on the classification of the Aplocheiloidei. Ichthyologi - infrequently flooded mangrove swamp. Northeast Gulf cal Exploration of Freshwaters , 15 : 105-120. Science , 10 : 63-68. Costa, W. J. E. M. 2004b. Kryptolebias , a substitute Taylor, S. 1990. Adaptive specializations of the cyprin - name for Cryptolebias Costa, 2004 and Kryptolebiati - odont fish Rivulus marmoratus . Biological Sciences , nae, a substitute name for Cryptolebiatinae Costa, 53 : 239-248. 2004 (Cyprinodontiformes: Rivulidae). Neotropical Taylor, W. R. & G. C. Van Dyke. 1985. Revised proce - Ichthyology , 2: 107-108. dures for staining and clearing small fishes and other Costa, W. J. E. M. 2004c. A new killifish genus and vertebrates for bone and cartilage study. Cybium , 9: species from the coastal plains of north-eastern Brazil 107-109. (Teleostei: Cyprinodontiformes: Rivulidae). Zootaxa , Turner, B. J., W. P. Davis & D. S. Taylor. 1992. Abun - 642 : 1-10. dant males in populations of a selfing hermaphroditic Davis, W. P., D. S., Taylor & B. J. Turner. 1990. Field fish, Rivulus marmoratus , from Belize cays. Journal of observations of the ecology and habits of mangrove Fish Biology , 40 : 307-310. rivulus ( Rivulus marmoratus ) in Belize and Florida Vermeulen, F. B. M. & T. Hrbek. 2005 Kryptolebias (Teleostei: Cyprinodontiformes: Rivulidae). Ichthyolog - sepia n. sp. (Actinopterygii: Cyprinodontiformes: Rivul - ical Exploration of Freshwaters , 1: 123-134. idae), a new killifish from the Tapanahony River Harrington, R. W. 1961. Oviparous hermaphroditic fish drainage in southeast Surinam. Zootaxa , 928 : 1-20. with internal self-fertilization. Science , 134 : 1749- 1750. aqua vol. 11 no. 1 - 2006 12 aqua, Journal of Ichthyology and Aquatic Biology

Centropyge abei , a new species of deep-dwelling angelfish (Pomacanthidae) from Sulawesi, Indonesia

1 2 3 Gerald R. Allen , Forrest Young and Patrick L. Colin

1) Western Australian Museum, Perth, Australia; mailing address: 1 Dreyer Road, Roleystone, WA 6111, Australia. 2) 10602 7 th Avenue Gulf, Marathon, Florida 33050, USA. 3) Coral Reef Research Foundation , Box 1765, Koror, PW 96940, Palau.

Accepted: 30.01.06 Keywords tige Farbgebung unterscheidet sich von allen anderen Taxonomy, marine fishes, Centropyge , new species, Angehörigen der Familie: eine ausgedehnte schwarze Pomacanthidae, Indonesia Fläche am Rücken und auf der Rückenflosse, das an der Seite in einen Gelbton übergeht, ein weißer Streifen Abstract hinter dem Kopf und weiße Schwanzflosse. A new species of pomacanthid fish, Centropyge abei , is described from a single specimen, 90.8 mm SL, col - Résumé lected from a depth of 120 m during deep diving opera - On décrit une nouvelle espèce de Pomacanthidé, Cen - tions off Manado, Sulawesi in Indonesia during April tropyge abei à partir d'un seul spécimen, d'une LS de 2005. It was also observed in similar habitat at Palau 90,8 mm, collecté à une profondeur de 120 m au cours between 110-155 m using a research submersible. de plongées profondes au large de Manado, Sulawesi, The combination of morphological features that en Indonésie, au mois d'avril 2005. Cette espèce a aussi include large scales (43-45 in lateral row from upper été observée, dans un habitat similaire, à Palau, entre opercle to caudal fin base), 3-4 preopercular spines, a 110 et 155 m, à l'aide d'un sous-marin de recherche. relatively narrow supracleithrum with pronounced ser - Une combinaison de caractéristiques morphologiques: rae on its posterior upper margin, serrate interopercu - de larges écailles (43 - 45 sur le rang horizontal du haut lum, and a posterior margin of the preorbital bone that is de l'opercule à la base de la caudale), 3 - 4 rayons durs attached and hidden by skin and scales serve to distin - préoperculaires, un supercleithrum relativement étroit guish it from all known angelfishes, although it appears avec des dentelures prononcées sur la marge su- to be a modified Centropyge . Moreover, its distinctive périeure postérieure, un interopercule dentelé et une colour pattern consisting of an extensive black area on marge postérieure de l'os préorbital qui est fixée et the back and dorsal fin that grades to yellow on the side, cachée par la peau et les écailles, aident à le distinguer white bar behind the head, and white caudal fin, is de tous les poissons-anges, même si elle ressemble à unlike that of any known species in the family. un Centropyge altéré. En outre, son patron de colora- tion caractéristique: une large zone noire sur le dos et Zusammenfassung la dorsale qui devient jaune sur le flanc, une barre blan- Beschrieben wird die neue Kaiserfischart Centropyge che derrière la tête et une caudale blanche, ne ressem - abei nach einem einzelnen Exemplar mit 90,8 mm SL, ble à aucun de ceux d'espèces connues dans la famille. das in einer Tiefe von 120 Metern bei Tieftauchgängen vor Monado, Sulawesi, in Indonesien im April 2005 Sommario gefangen wurde. Weitere Exemplare konnten in einem Una nuova specie di pomacantide, Centropyge abei , ähnlichen Habitat bei Palau in einer Tiefe von 110 bis è descritta sulla base di un singolo esemplare di 90.8 155 Metern von einem Forschungstauchboot aus beo- mm SL, raccolto a 120 m di profondità durante immer - bachtet werden. Große Schuppen (43-45 in einer seitli- sioni profonde al largo di Manado, Sulawesi in Indone - chen Reihe vom oberen Operculum bis zur Schwan- sia nell’Aprile 2005. E’ stato anche osservato in un zflossenbasis), 3-4 Präoperculum-Stacheln, ein relativ habitat simile a Palau a profondità di 110-155 m utiliz - schmales Supracleithrum mit deutlichen Sägezähnchen zando un sottomarino da ricerca. La combinazione di am hinteren oberen Rand, ein gesägtes Interoperculum tratti morfologici che includono larghe scaglie (43-45 in sowie ein Hinterrand des Präorbitalknochens, den linea laterale dall’opercolo superiore alla base della pin- anheftende Haut und Schuppen verdecken – das sind na caudale), 3-4 spine preopercolari, un supracleitrum Merkmale, die insgesamt diese neue Art von allen relativamente stretto con pronunciate dentellature sul anderen Kaiserfischen unterscheiden, wobei sie schon suo margine superiore, interopercolo seghettato, e als zu Centropyge gehörig erscheint. Auch die einzigar - margine posteriore dell’osso preopercolare attaccato e

13 aqua vol. 11 no. 1 - 2006 Centropyge abei , a new species of deep-dwelling angelfish (Pomacanthidae) from Sulawesi, Indonesia nascosto dalla pelle e dalle scaglie, servono a distin - Centropyge abei n. sp. guerlo da tutti glia altri pesci angelo, sebbene appaia un (Figs 1-2) Centropyge modificato. Inoltre, la distinta colorazione consistente in un’estesa area nera sul dorso e sulla Holotype: NCIP 6305, 90.8 mm SL, south-east pinna dorsale che degrada nel giallo sui fianchi, una side of Manado Tua (approximately 01°6.2040’S, barra bianca dietro la testa e una pinna caudale bianca 124°7.1222’E), Sulawesi, Indonesia, 120 m, collected risulta diversa da tutte le specie note della famiglia. with quinaldine and hand net by F. Young, 27 April 2005. Introduction Angelfishes of the family Pomacanthidae are among Diagnosis the most conspicuous inhabitants of coral reefs, occur - A species of the pomacanthid genus Centropyge with ring in both shallow and relatively deep water. Allen et the following combination of characters: dorsal rays al. (1998) reviewed the family, recognizing 83 species XIII,17; anal rays III,18; pectoral rays 16; scales rela - worldwide, including 70 from the Indo-Pacific region. A tively large, 43-45 in lateral row from upper end of gill more recent (2003) treatment of the group by Debelius opening to base of caudal fin; lateral line with 37 scales, et al . recognized 75 Indo-Pacific species, including 28 terminating below end of dorsal fin, but re-appearing on Indo-Pacific members of the genus Centropyge Kemp, 7 scales of caudal peduncle; gill rakers on first arch 5 + 1860. 12; maximum body depth 1.6 in SL; preopercular spines The present paper describes a new species of deep- 3 or 4; length of primary preopercular spine 2.0 in head dwelling angelfish that is provisionally assigned to the length; teeth tricuspid, about 50-60 in outer row in each genus Centropyge . It was first sighted during subme- jaw; exposed margins of preorbital, interopercle, preop - rsible dives along the outer slope of Palau in 2001 by the ercle, subopercle, supracleithrum, and posttemporal third author. A single specimen, described herein, was bones serrate; posterior margin of preorbital bone subsequently collected during a series of deep re- attached and hidden by skin and scales; live coloration breather scuba dives by the second author off Manado, yellow with black upper third of back, dorsal fin, and Sulawesi in Indonesia during April 2005. Although the upper part of head. A broad white bar immediately ante - primary goal of this expedition, to locate and film a re- rior to dorsal fin origin, extending to at least level of cently described coelacanth, was unsuccessful, several upper opercular margin. Caudal fin and peduncle white. valuable collections of small reef fishes were obtained. Description Materials and Methods Dorsal rays XIII,17 (last divided to base); anal rays The specimen of the new angelfish was obtained with III,18 (last divided to base); pectoral rays 16 (upper two the aid of mixed gas, re-breather scuba equipment and lowermost unbranched); pelvic rays I,5; transverse allowing dives to working depths of approximately 150 scale rows from upper end of gill opening to base of m. A mild anaesthetic, quinaldine sulphate, was used to caudal fin 43-45; scale rows above lateral line to origin partially narcotise fish specimens prior to capture with a of dorsal fin 9; scale rows below lateral line to origin of hand net. anal fin about 20; lateral line in two sections, the ante - Standard length (SL) was measured from the front of rior part extending from upper edge of gill cover to below the upper lip to the base of the caudal fin (end of hypural level of posteriormost dorsal fin ray containing 37 scales plate). Total length (TL) was measured from the front of and the posterior part containing 7 scales along middle the upper lip to the tip of the longest caudal fin ray. The head length was taken from the front of the upper lip to the end of the opercular membrane. The depth was measured just in front of the anal fin to the extreme base of the dorsal spines. The length of the caudal peduncle was measured horizontally from a vertical at the rear base of the anal fin to the caudal fin base. Measure - ments of the dorsal and anal spines and soft rays were made from the distal tips to the extreme bases of these elements (aided by x-ray). A radiograph was employed to examine various osteological features. The upper limb gill raker count is given first; the raker at the angle is included in the lower limb count. Proportional measurements expressed in thousandths of the standard length are provided in parenthesis in the description that appears below. The holotype – and only Fig. 1. Freshly collected specimens of Centropyge abei known specimen - is deposited at Pusat Penelitian dan n. sp., holotype, 90.8 mm SL, Manado Tua, Sulawesi, Pengembangan Oseanologi, Jakarta, Indonesia (NCIP). Indonesia. Photo by Kotaro Yoshimura. aqua vol. 11 no. 1 - 2006 14 Gerald R. Allen, Forrest Young and Patrick L. Colin of caudal peduncle; transverse scale rows on opercle 7; Upper and lower lips about equal in size, broadly scaled gill rakers on first branchial arch 5 + 12, total rakers 17, except anterior edges, their maximum width contained the rakers short and stubby, much shorter than gill fila - 2.8 in diameter of orbit. Teeth slender, elongate (the ments; vertebrae 10 + 14. longest 4.6 in orbit), close-set, flexible, tricuspid (large Body ovate, the depth 1.6 (61.3 % of SL) in SL, and central cusp notably longer than smaller lateral ones), in compressed, the maximum width 2.6 (21.4 % of SL) in 5-6 rows in each jaw, about 50-60 in outer row of each depth; head length 3.2 (30.9 % of SL) in SL; dorsal pro - jaw. No teeth on roof of mouth. Tongue short and file of forehead relatively steep and straight, forming an broadly rounded. angle of about 45 degrees to the horizontal; snout 2.6 Nostrils anterior to centre of eye, the posterior opening (12.1 % of SL) in head length, diameter of orbit 3.1 (10.0 relatively large and ovate with thin, slightly raised rims; % of SL) in head length; interorbital space slightly con - anterior nasal opening about one-fourth size of posterior vex, the least bony width 3.5 (8.8 % of SL) in head one, in a membranous tube with a posterodorsal flap; length; caudal peduncle much deeper than long, the Most head pores inconspicuous (due to dense covering least depth 2.2 (14.3 % of SL) in head length; length of of tiny scales) except for prominent pore just anterior to caudal peduncle 6.0 (5.2 % of SL), in head length. anterior nasal opening. Gill membranes narrowly Mouth relatively small, terminal, the gape forming an attached to isthmus. Longest gill filament on first arch angle of about 30 degrees to the horizontal, the maxilla contained 1.9 times in orbit. Gill rakers short, the longest reaching a vertical at about front of anterior nostril. 15.1 in orbit.

Fig. 2. Close-up photograph of head of preserved holotype of Centropyge abei n. sp. Photo by G. Allen.

15 aqua vol. 11 no. 1 - 2006 Centropyge abei , a new species of deep-dwelling angelfish (Pomacanthidae) from Sulawesi, Indonesia

Upper edge of opercle obtusely rounded, without a incised; caudal fin scaled nearly to posterior margin; spine. A prominent large spine at corner of preopercle, pectoral and pelvic fins densely scaled at base, with tiny longer than orbit, the spine length (measured along scales extending on surface of rays (but not on mem - lower edge) contained 2.0 (15.3 % of SL) times in head branes) nearly to posterior margin. length; a series of 2 (right side) or 3 smaller preopercle Lateral line relatively inconspicuous, its presence indi - spines immediately below and anterior to primary spine, cated by smaller scales (including tiny auxiliary scales), the largest (posteriormost) one-fourth (right side) to one- gently arching across back, originating at upper corner half length of primary spine; margin of preopercle with of gill opening and terminating below end of soft dorsal 23-25 serrae; anterior and ventral margins of preorbital fin; additionally, 7 tubed scales midlaterally on caudal with 5-9 serrae, the posterior margin attached and hid - peduncle. den by skin and scales; lower exposed margin of Caudal fin rounded, its length 1.2 (25.3 % of SL) in interopercle with 2-5 serrae; margin of subopercle with head length. Origin of dorsal fin slightly anterior to a ver - 12-13 serrae; exposed posterior margin of supraclei - tical at upper end of gill opening. Dorsal spines pro - thrum relatively narrow and covered with scales, its gressively longer to last spine; first dorsal spine 2.7 upper posterior edge with 10-13 serrae; posterior edge (11.5 % of SL), second dorsal spine 2.1 (14.5 % of SL), of posttemporal with 4-7 serrae. third dorsal spine 1.9 (16.0 % of SL), all in head length; Scales of body more or less arranged in regular trans - membranes between first eight dorsal spines and three verse rows and coarsely ctenoid (up to 25 ctenii on anal spines deeply incised; posterior margin of soft por - exposed posterior margins); auxiliary scales mainly tions of dorsal and anal fins angular, the longest dorsal confined to lateral line and anteriormost portion of body; and anal rays reaching as far posterior as level of mid - head fully scaled except anterior edges of lips; dorsal dle of caudal fin, their length 1.5 (21.0 % of SL) and 1.4 and anal fins scaled nearly to margins except anteriorly (22.8 % of SL) respectively in head length; origin of anal in spinous portion where fin membranes are deeply fin below base of ninth or tenth dorsal spine; first anal

Fig. 3. Underwater photographs of Centropyge abei n. sp., showing different swimming postures ( A-C ), approximately 12 cm TL, taken from Deepworker submersible at Palau in about 120 m depth. Photo by Coral Reef Research Foundation. aqua vol. 11 no. 1 - 2006 16 Gerald R. Allen, Forrest Young and Patrick L. Colin spine 1.8 (15.5 % of SL), second anal spine 1.5 (20.7 % small and serrate, remote from suboperculum; hindmar - of SL), third anal spine 1.4 (21.4 % of SL), all in head gin of preorbital free, prominently serrate or with strong length; pectoral fins relatively short and moderately spines; interorbital width equal to or less than eye; pointed, reaching a vertical about equal to anal opening, scales on operculum in five or fewer transverse rows. their length 1.0 (30.2 % of SL) in head length; pelvic fin The salient features of the new species include the fol - tips filamentous, nearly reaching to anal fin origin, their lowing features that agree with the definition of Cen - length 1.1 (28.9 % of SL) in head length; pelvic fin spine tropyge: a relatively small body size, small scales 1.5 (20.6 % of SL). arranged in regular transverse rows, a relatively small, Colour when fresh (Fig. 1): lower two-thirds of body serrate interoperculum and the least interorbital width yellowish, grading to yellow-white ventrally; upper third slightly narrower than eye diameter. However it differs of body dusky brown quickly grading to jet black, includ - from Centropyge in having an attached posterior margin ing dorsal fin; chalky white bar immediately anterior to of the preorbital and by its possession of seven trans - dorsal fin origin, extending to level of uppermost opercle verse scale rows on the operculum. In addition, the margin, its width equal to 1.0-1.3 eye diameter; upper colour pattern, particularly the broad white bar behind portion of cheek charcoal, grading to black on upper the head, is a feature that is absent in all other members part of head; lips, snout tip, lower part of head, interop - of the genus. This marking is most prevalent in the ercle, and opercle margin greyish white; opercle greyish genus Chaetodontoplus , which is also distinguished by white with black margin, most pronounced above pec - very small scales (>85), spineless interoperculum, a toral fin base; pectoral, pelvic, and anal fins yellow; cau - broad interorbital that exceeds the eye diameter, and dal fin and peduncle white. larger body size (usually >15 cm SL). Photos of live individuals taken from the submersible Apolemichthys is the only other generic possibility. in Palau indicate a similar coloration with the following Indeed, our first impression when viewing both the live exceptions: 1. the snout and lips are paler (whitish); 2. fish underwater and photographs was that the new the prominent white bar behind the head is broader and species belonged to this genus. The eight known mem - extends farther ventrally (to lower margin of preopercle); bers of this genus (Allen et al. 1998) have the same and 3. the dark pigmentation on the upper body is far general body shape as the new species, similar sized more extensive, covering the entire upper half. scales and at least some of the species are known to Colour in alcohol : most of body and lowermost part frequent deep reefs in excess of 100 m. However, the of head pale tannish; upper third of body and adjacent new species differs in several key features that appear dorsal fin black, the coloration most intense on dorsal to preclude its inclusion, particularly the serrate interop - fin; head with dusky grey cheek, grading to black or erculum (smooth in Apolemichthys) and the multispined charcoal on upper part of head; lips, interopercle, pre - preoperculum (only one spine in Apolemichthys). In opercle margin pale tan; opercle mainly pale tan with addition, radiograph evaluation of the new species indi - intense black margin on portion above pectoral fin base; cates that it lacks two features which, according to distinctive white bar immediately anterior to dorsal fin Heemstra (1984: Figs. 1 and 3), appear to be uniquely origin, extending to level of uppermost opercle margin, derived homologous characters that may serve to dis - its width about equal to eye diameter dorsally and about tinguish Apolemichthys from related genera: namely, 1.3 eye diameter ventrally; all fins except dorsal yellow - lateral expansion of the anterior 2-5 haemal spines ish tan. (unmodified in C. abei) and a supracleithrum that is ovate or oblong with its rear edge exposed (relatively Remarks narrow with rear edge covered with large scales in C. The new species appears to be most closely related to abei as in other pomacanthids). the genus Centropyge to which it is provisionally assigned. However, it possesses several morphological Ecological observations peculiarities (discussed below) that are indicative of The type locality was situated in 120 m depth at the possible separate subgeneric status. We hesitate to base of a 45 degree talus slope, on a general slope of describe it as a new subgenus in view of the current 60-80 degrees that was occasionally interrupted by rel - study of the familial classification by Richard Pyle of the atively flat plateaux. The bottom was primarily com - Bishop Museum in Hawaii. Our specimen is currently posed of variably sized rubble. The water temperature being studied by Pyle and its ultimate generic status will was 20-21 degrees C. The only other angelfishes seen be discussed by him in a future publication. (but not collected) at this depth were an unknown and The genus Centropyge has been traditionally defined possibly new species of Genicanthus , and Centropyge (see Weber and De Beaufort, 1936: 159; Fraser-Bruner, tibicen , normally a shallow reef dweller, which was com - 1933) as relatively small angelfishes (usually < 10 cm mon between 100-110 m. SL) with the following combination of features: scales The new species was seen on eight out of 40 sub - large, usually 50 or less in lateral row between upper mersible dives (Deepworker 2000) on the outer slope of operculum and caudal fin base, arranged in more or Palau in 2001. Observations ranged between 110 and less regular transverse rows; interoperculum relatively 155 m (365-515 ft) with water temperatures 17-22°C.

17 aqua vol. 11 no. 1 - 2006 Centropyge abei , a new species of deep-dwelling angelfish (Pomacanthidae) from Sulawesi, Indonesia

The region is prone to rapid thermal changes of several Acknowledgements degrees C (Wolanski et al . 2002). The fish was immedi - We thank Masa Iwata for organizing the expedition ately recognized as a new species on the basis of its and providing financial support, Dr. Kasim Moosa for distinct colour pattern, but since the submarine was not arranging for collecting permits and providing his wis - equipped to collect fishes, no attempt was made to cap - dom and guidance. Ben Daughtry capably assisted on ture a specimen. Still photos (and video footage) were all the deep dives and assumed responsibility for the obtained of the new fish, one of which is included as technical aspects, enabling Forrest Young to concen - Fig. 3. trate on fish collections. Captain Billy Deans managed Centropyge abei was not common at Palau. It was surface operations and various aspects of diving safety. seen on only eight of 40 dives within the depth range Heath Laetari, Kenichi Fuji, Kotaro Yoshimura and Larry indicated in the previous paragraph. Never more than Wright provided backup safety for the deep-diving team. one pair was seen on a single dive, and it is likely that Special thanks are due to Mark Erdmann for guiding us some of the observations on different days involved the to suitable dive sites including potential coelacanth loca - same fish, as the dives were often made in the same tions. Danny Charlton provided his dive facility that we locations. Either solitary individuals or pairs were used for an operational base. observed. Apolemichthys trimaculatus was the only Observations and photographs of the new angelfish other pomacanthid seen at similar depths (110-125 m). were made possible in Palau through the charter of the The Palauan habitat consisted of a very steep (60-80 Deepworker 2000 for activities associated with the US degree) limestone slope with accumulations of reef National Cancer Institute’s shallow water marine collec - limestone talus below about 120 m, forming small over - tion and taxonomy contract (N02-CM-77249) to the hangs and crevices. The fish often sought shelter in Coral Reef Research Foundation. All CRRF sub - such areas when the sub first approached, but would mersible pilots (P. L. Colin, L. J. Bell, M. N. Dawson and often re-emerge quickly and did not seem greatly dis - L. E. Martin) contributed observations on the occur - turbed by the presence of the sub and its lights within rence of this new fish. several meters. However, they fled to shelter when Glenn Moore of the Western Australian Museum approached at closer range. The water was very clear kindly provided a radiograph of the holotype of C. abei . and the ambient light was strong enough to clearly see Finally, we thank John E. Randall and Richard Pyle for the major substratum features, but it was quite dark critically reviewing the manuscript. beneath overhangs and in crevices. Currents were not strong in the areas, which had to be relatively sheltered References in order for the submersible to operate safely. Allen, G. R., Steene, R. & M. Allen. 1998. Guide to The type locality (near Manado, Sulawesi) and Palau Angelfishes and Butterflyfishes. Odyssey Press, San are separated by a distance of approximately 1,235 km. Diego. It likely C. abei is much more widespread, but due to the Debelius H., Tanaka, H. & R. H.Kuiter. 2003. difficulties of sampling deep reef environments, its pre - Angelfishes, A comprehensive Guide to Pomacan- cise distribution remains unknown. The fish fauna of the thidae . TMC, Chorleywood, UK. deep reef or “twilight zone”, lying between about 70-200 Fraser-Brunner, A. 1933. A revision of the chaetodont m is very poorly documented and holds promise for fishes of the subfamily Pomacanthinae. Proceedings future discoveries. Limited observations suggest that of the Zoological Society of London 1933 (part 3, no. numerous shallow water species penetrate to consider - 30): 543-599. able depth, but the fauna also contains a wealth of Heemstra, P. C. 1984. Apolemichthys kingi , a new undescribed deep-dwelling reef fishes. species of angelfish (Pomacanthidae) from South Africa, with comments on the classification of Etymology angelfishes and a checklist of the pomacanthids of the The species is named abei in honour of Dr.Yoshitaka western Indian Ocean. J. L. B. Smith Institute of Abe, without whose faith, guidance and support, none of Ichthyology Special Publication No. 35 : 1-17. the work would have been possible. Dr. Abe is the direc - Weber, M. & L. F. de Beaufort . 1936. The fishes of the tor of Aquamarine Fukushima, a world class public Indo-Australian Archipelago. VII. Perciformes (contin - aquarium in Fukushima Prefecture, Japan. He is largely ued) families: Chaetodontidae, Toxotidae, Mon - responsible for many innovations in aquarium science odactylidae, Pempheridae, Kyphopsidae, Lutjanidae, and design including display of large tunas up to 100kg, Lobotidae, Sparidae, Nandidae, Sciaenidae, Malacan - jellyfish keeping, and the first public display of large thidae, Cepolidae. E. J. Brill Ltd., Leiden. Fish. Indo- hammerhead sharks. Aquamarine Fukushima and Dr. Aust. Arch. v. 7: i-xvi + 1-607. Abe provided the entire budget and material support for Wolanski, E., P. L. Colin, J. Naithani, E. Deleersni - the deepwater operations that resulted in the collection jder & Y. Golbuu. 2004. Large amplitude, leaky, of the new angelfish. island-generated, internal waves around Palau, Micronesia. Estuarine, Coastal and Shelf Science, 60 : 705-716. aqua vol. 11 no. 1 - 2006 18 aqua, Journal of Ichthyology and Aquatic Biology

Amblyeleotris neumanni , a new species of shrimp goby from New Britain

John E. Randall and John L. Earle

Bishop Museum, 1525 Bernice St., Honolulu, HI 96817-2704, USA

Accepted: 30.01.06

Keywords de l'île Lolobau, Nouvelle Angleterre, Papouasi Nouvelle Taxonomy, Gobiidae, Amblyeleotris , new species, Guinée. Il a été observé en association symbiotique New Britain avec les crevettes Alpheus rapacida et A. sp., avec lesquelles il partage un terrier. Il se caractérise par la Abstract présence de 12 ou 13 rayons mous à l'anale, 19 rayons The gobiid fish Amblyeleotris neumanni is described pectoraux, 98 - 100 écailles en rangée longitudinale, les as a new species from three specimens, 44.2-53.9 mâles, avec les 3e et 5e rayons filamenteux, une cau - mm SL, collected in 26 m off Lolobau Island, New dale lancéolée, des nageoires pelviennes complète - Britain, Papua New Guinea. It was observed in sym - ment soudées, un frenum pelvien, et par la coloration: biotic association with the snapping shrimps Alpheus quatre larges barres brun orange sur le corps entre - rapacida and A. sp., with which it shares a burrow. It coupées de taches foncées irrégulières, des ocelles is distinctive in having 12 or 13 dorsal soft rays, 13 orange sur les rayons durs et des rayons mous des dor - anal soft rays, 19 pectoral rays, 98-100 scales in lon - sales, la base de l'anale marquée longitudinalement de gitudinal series, males with filamentous third to fifth lignes étroites adjacentes, noire, orange, bleue et bleu- spines, lanceolate caudal fin, pelvic fins fully joined, vert; pelviennes bleu pâle ou verte s. pelvic frenum, and in colour: four broad dark orangish brown bars on body with irregular dark markings Sommario between; ocellated orange spots on spines and rays Il gobide Amblyeleotris neumanni è descritto come of dorsal fins; basal part of anal fin with adjacent nar - specie nuova sulla base di tre esemplari di 44.2-53.9 row longitudinal bands of black, orange, blue, and mm SL, raccolti a 26 m al largo dell’isola di Lolobau, blue-green; pelvic fins pale blue or green. Nuova Britannia, Papua Nuova Guinea. E’ stato osser - vato in associazioni simbiotiche con il gamberetto Zusammenfassung Alpheus rapacida e A. sp., con cui condivide la tana. Si Beschrieben wird als neue Art die Meergrundel distingue epr avere 12 o 13 raggi molli dorsali, 13 raggi Amblyeleotris neumanni nach drei Exemplaren mit 44,2- molli anali, raggi 19 pettoral, 98-100 scaglie in serie lon - 53,9 mm SL, die in 26 Metern Tiefe vor den Lolobau- gitudinale, i maschi con terza, quarta e quinta spina fila - Inseln, New Britain, Papua-Neuguinea gefangen wur - mentosa, pinna caudale lanceolata, pinne pelvicche den. Sie wurden in Symbiose mit dem Knallkrebs Alphe- completamente unite, frenum pelvico presente e la colo - us rapacida und einer weiteren Art beobachtet, mit de- razione: quattro ampie barre arancio-marrone scuro sul nen sie die Höhle teilten. Ihre Unterscheidungsmerk - corpo con macchie irregolari inframmezzate; macchie male sind 12 oder 13 weiche Rückenflossenstrahlen, 13 arancio ocellate sulle spine e i raggi della dorsale; base weiche Analflossenstrahlen, 19 Brustflossenstrahlen, 98 della pinna anale con strette bande ravvicinate longitu - - 100 Schuppen in Längsreihen, bei den Männchen dinali di colore nero, arancio, blu e grigio-blu; pinne pel - fädige Stacheln an 3. und 5. Stelle, eine lanzettförmige viche blu pallido o verde. Schwanzflosse, vollständig vereinte Bauchflossen, ein Beckenband und die Farbgebung: vier breite, dunkle Introduction orangebraune Bänder auf dem Rumpf mit unregelmäßi - Species of 13 genera of gobiid fishes live in symbi - gen dunklen Markierungen dazwischen; orangefarbene otic association with snapping shrimps of the genus Augenflecken auf Stacheln und Flossenstrahlen der Alpheus . The shrimp builds and maintain a burrow in Rückenflossen; am basalen Teil der Analflosse schmale sedimentary substrata. The goby, with its superior Längsstreifen in Schwarz, Orange, Blau und Blaugrün; vision and cephalic sensory system, serves as the Bauchflossen blassblau oder -grün. sentinel at the burrow entrance, quickly taking refuge in the burrow, if needed. The most speciose of the 13 Résumé genera is the Indo-Pacific genus Amblyeleotris , which Le gobie Amblyeleotris neumanni est décrit en tant includes some of the most colourful fishes of our qu'espèce nouvelle sur base de trois spécimens, d'une planet. It was described by Bleeker (1874), designat - LS de 44,2 à 53,9 mm, et a été collecté, à 26 m, au large ing Eleotris periophthalmus Bleeker as the type

19 aqua vol. 11 no. 1 - 2006 Amblyeleotris neumanni , a new species of shrimp goby from New Britain species. In a paper describing five new species of Meristic and morphometric data in parentheses refer Amblyeleotris from islands of Oceania, Randall (2005) to paratypes. provided a diagnosis of the genus and a brief histori - cal resume of the species. A new species from the island of New Britain, Papua New Guinea is described Amblyeleotris neumanni n. sp. in the present paper, bringing the total number of (Figs. 1-4; Table I) species in the genus to 34. The opportunity to collect and photograph fishes in Holotype: BPBM 39048, 54.9 mm, Papua New New Britain was provided by a cruise on the dive ves - Guinea, New Britain, Lolobau Island, east end off red sel Pelagian , which commenced in July, 2002 at cliff, 4°54.1’S, 151°12.7’E, dark silty sand and rubble, Kimbe Bay on the north coast and sailed east to 26 m, collected with spear by J. E. Randall, 3 August Rabaul. About half way to Rabaul we stopped to dive 2002. at Lolobau Island, an active volcano 6.5 by 10 km in Paratypes: BPBM 39049, 44.2 mm and USNM size and 832 m high. We were joined on the dive by 385742, 46.3 mm, same data as holotype. Mike Neumann who had financed the cruise. We spotted a new species of Amblyeleotris on a bottom of Diagnosis dark brown sand and rubble at a depth of 26 m. Dorsal rays VI+I,12-13; anal rays I,13; pectoral rays Underwater photographs were taken, and three spec - 19; longitudinal scale series 98-100; no median pre - imens were collected with a small multiprong Hawai - dorsal scales; scales on side of nape extending for - ian sling spear. ward to above middle of opercle; body depth 5.5-5.8 in SL; head length 3.45-3.5 in SL; snout length 1.75 in Materials and Methods HL; gill opening extending forward to above middle of The holotype and one paratype of the new species peopercle; third to fifth dorsal spines of male pro - are deposited in the Bishop Museum, Honolulu longed as filaments; caudal fin lanceolate, 2.6-2.85 in (BPBM), and one paratype in the U.S. National SL; pelvic fins fully joined medially, reaching origin of Museum of Natural History, Washington, D.C. anal fin; pelvic frenum present; four orangish brown (USNM). bars on body with irregular dark markings between; The length of specimens is given as standard length rays of dorsal fins with ocellated orange spots, the (SL), measured from the median anterior end of the soft rays tipped with orange; basal part of anal fin with upper lip to the base of the caudal fin (posterior end of narrow adjacent bands of black, orange, blue, and the hypural plate); body depth from the origin of pelvic blue-green; pelvic fins pale blue or green. fins, and body width at the origin of the pectoral fins; head length (HL) from the upper lip to the posterior Description end of the opercular membrane, and head width over Dorsal rays VI+I,13 (12-13); anal rays I,13; all dor - the posterior margin of the preopercle; orbit diameter sal and anal soft rays branched, the last to base; pec - is the greatest fleshy diameter, and interorbital width toral rays 19, the uppermost and lower 1 or 2 un- the least bony width; snout length is measured from branched; pelvic rays I,5, all soft rays branched, the the median anterior point of the upper lip to the near - medial rays fully united by membrane; pelvic frenum est fleshy edge of the orbit; upper jaw length from the present; segmented caudal rays 17, 14 (13-14) bran- same anterior point to the posterior end of the maxilla; ched; upper and lower unsegmented caudal rays 7; caudal peduncle depth is the least depth, and caudal longitudinal scale series 98 (99-100); transverse scale peduncle length the horizontal distance between ver - rows 31 (32-33); circumpeduncular scales about 25; ticals at the rear base of the anal fin and the caudal fin gill rakers 4+13 (4+10-11); pseudobranch with 8 base; lengths of spines and rays are measured to fleshy lobes; branchiostegal rays 5; vertebrae 25. their extreme bases; caudal fin and pectoral fin Body depth 5.8 (5.5-5.8) in SL; body width 1.75 lengths are the length of the longest ray; pelvic fin (1.65-1.8) in body depth; HL 3.5 (3.45-3.5) in SL; head length is measured from the base of the pelvic spine width 2.5 (2.4-2.45) in HL; snout length 1.75 (1.75) in to the tip of the longest soft ray. Morphometric data HL; dorsal profile of snout moderately steep, forming are presented in Table I as percentages of the stan - an angle of about 40° to horizontal axis of head and dard length. Proportional measurements in the text body; orbit diameter 3.95 (3.75-3.8) in HL; upper edge are rounded to the nearest 0.05. of eye extending above dorsal profile of head; interor - The count of scales in longitudinal series is made bital space extremely narrow, the least width 36 (41) in from above the upper end of the gill opening to the HL; caudal peduncle depth 3.0 (3.1-3.15) in HL; cau - base of the caudal fin; scales in transverse series are dal peduncle length 1.75 (1.7-1.75) in HL. counted from the origin of the anal fin obliquely Mouth oblique, forming an angle of about 35° to hor - upward to the base of the first dorsal fin; the count of izontal axis of head and body, with the lower jaw gill rakers is made on the first gill arch, those on the slightly projecting; maxilla reaching to or slightly pos - upper limb given first. terior to a vertical through centre of eye, the upper jaw aqua vol. 11 no. 1 - 2006 20 John E. Randall and John L. Earle length 2.6 (2.4-2.55) in HL; front of upper jaw with two Scales small for the genus, progressively smaller well-spaced, incurved canine teeth, the symphyseal anteriorly, extending on side of nape to above middle gap in holotype nearly equal to pupil diameter, the of opercle, but none in median predorsal zone or length of the largest and more medial of the canines prepectoral area (if embedded scales are present in two-thirds pupil diameter in length; side of upper jaw these two apparently naked areas, none could be dis - with a row of 18 slender incurved canine teeth in holo - lodged); scales ctenoid posteriorly on body, becoming type, progressively shorter posteriorly; an inner band cycloid anterior to origin of second dorsal fin; small of small, strongly incurved villiform teeth in three or cycloid scales present on chest; no scales on fins four rows at front of jaws, narrowing to a single row except basal fourth to fifth of caudal fin. posteriorly; front of lower jaw with three pairs of Origin of first dorsal fin over inner base of pelvic fins, incurved slender canine teeth smaller than those of the predorsal length 3.4 (3.3) in SL; first dorsal spine upper jaw and an inner band of incurved villiform teeth 1.8 (1.7-2.05) in HL; fourth and fifth dorsal spines in at most three rows; side of lower jaw nearly halfway longest and subequal, 1.2 (1.2 in male paratype and from symphysis with three large close-set retrorse 1.6 in female paratype) in HL; third to fifth dorsal canine teeth, the largest as long as largest canine at spines prolonged as short filaments in males; spine of front of upper jaw; remainder of lower jaw with a sin - second dorsal fin 2.2 (1.95-2.l) in HL; penultimate and gle row of close-spaced slender teeth smaller than preceding dorsal soft rays subequal and longest, 1.55 those of upper jaw; no teeth on vomer or palatines; (1.4-1.65) in HL; origin of anal fin a little anterior to tongue broadly rounded; inner edge of lips papillose. base of second dorsal soft ray, preanal length 1.8 Gill opening extending forward to below middle of (1.8) in SL; anal spine 3.45 (3.45-3.85) in HL; penulti - preopercle (about one-half eye diameter posterior to mate and preceding anal soft rays longest, 1.5 (1.6) in eye). Gill rakers short, the longest about half length of HL; caudal fin lanceolate, 2.6 (2.6-2.85) in SL; pec - longest gill filaments. toral rays pointed, the middle rays longest, 1.15 (1.15- Posterior nostril anterior to lower third of eye just 1.2) in HL; prepelvic length 3.4 (3.25-3.5) in SL; pelvic before fleshy edge of orbit, the aperture small, only spine 2.65 (2.7-3.05) in HL; fifth pelvic soft ray slightly larger than sensory pores of head, with a very longest, extending to origin of anal fin, 3.7 (3.7-3.85) low rim; anterior nostril ventroanterior to posterior in SL; pelvic frenum well developed, the membrane nostril, at end of a short membranous tube, nearly reaching about two-thirds length of pelvic spine. reaching edge of snout when depressed forward. Colour of male holotype in life: shown in Fig. 1. A Pores of cephalic sensory system and sensory papil - smaller male is illustrated in Fig. 2, and females in lae on cheek and opercle as shown in illustration of Figs. 3 and 4. Noteworthy colour features are the four Amblyeleotris japonica by Akihito in Masuda et al. dark orangish brown bars on the body with irregular (1984: 255, fig. 95). dark markings in the bluish to whitish interspaces, the

Fig. 1. Holotype of Amblyeleotris neumanni n. sp., male, BPBM 39048, 54.9 mm, Lolobau Island, New Britain. Photo by M. Neumann.

21 aqua vol. 11 no. 1 - 2006 Amblyeleotris neumanni , a new species of shrimp goby from New Britain

Table I. Proportional measurements of type specimens dark brown, blue, and orange markings on the head, of Amblyeleotrus neumanni n. sp. as percentages of the standard length. the ocellated orange spots on the dorsal spines (the spines of females tipped with orange), the second Holotype Paratypes dorsal fin with an orange spot just posterior to ray tips, the basal part of anal with adjacent longitudinal bands BPBM BPBM USNM 39048 39049 385742 of black, orange, blue, and blue-green, the orange upper margin of the caudal fin, and the pale blue Sex male female male pelvic fins. Standard length (mm) 54.9 44.2 46.3 : pale tan, becoming Body depth 17.2 18.1 17.3 Colour of holotype in alcohol Body width 9.8 10.0 10.4 still paler ventrally, with four broad brown bars nar - Head length 28.4 28.7 28.9 rower than pale interspaces, the first on nape narrow - Head width 11.8 11.7 12.1 ing ventrally onto opercle, and the last on caudal Snout length 6.7 6.6 6.9 peduncle; pale interspaces with irregular narrow verti - Orbit diameter 7.2 7.5 7.7 cal dark bands dorsally on body, some commencing Interorbital width 0.8 0.7 0.7 on back in a distinct brown spot; interorbital space Upper jaw length 11.0 11.3 11.9 and upper edge of eyes dark brown, the pigment fol - Caudal peduncle depth 9.5 9.3 9.2 lowing sensory canal behind upper part of eye and Caudal peduncle length 16.3 16.3 16.7 continuing horizontally as a band to upper pore of pre - Predorsal length 29.7 30.3 30.4 opercular sensory canal; front of snout and medial Preanal length 55.0 55.7 56.2 Prepelvic length 29.2 30.9 28.5 part of lips and chin brown; a faint brown bar from Base of dorsal fins 55.1 54.5 53.3 below centre of eye to behind posterior end of upper First dorsal spine 15.5 14.9 16.8 jaw; ventral margin of gill membranes broadly dark Longest dorsal spine 23.5 18.3 24.4 brown; dorsal fins pale yellowish, the first dorsal Spine of second dorsal fin 13.1 13.5 14.8 dusky with a pale spots (fin damaged on holotype; Longest dorsal ray 18.3 17.4 20.6 spotting more evident on fin of male paratype); caudal Base of anal fin 29.7 27.2 28.0 fin pale, a little dusky on scaled basal part, with a dark Anal spine 8.2 7.5 8.4 brown streak on outer part of membrane between Longest anal ray 19.2 18.0 broken tenth and eleventh branched rays; anal fin transparent Caudal fin length 38.1 35.1 38.4 with a narrow, very dark brown stripe in lower middle Pectoral fin length 24.2 24.2 24.9 Pelvic spine length 10.7 9.4 10.6 part of fin, bordered above by a light brown band that Pelvic fin length 27.2 26.1 27.2 is bisected by a darker brown line; pectoral fins pale, the rays edged with light brown, with a dusky spot on

Fig . 2. Amblyeleotris neumanni n. sp., male, with Alpheus sp., Lolobau Island, New Britain. Photo by J. E. Randall. aqua vol. 11 no. 1 - 2006 22 John E. Randall and John L. Earle upper base; pelvic fins with brown membranes that Remarks are darker medially. This species was observed and collected only at one site in 26 m at Lolobau Island, New Britain. Figures 2 Etymology and 3 show its symbiotic partner Alpheus sp., proba - We are pleased to name this colourful goby in hon - bly an undescribed species related to A. brevirostris our of Mike Neumann, fellow diver, underwater pho - (Arthur Anker, pers. comm.). A second snapping tographer and good friend. shrimp, A. rapacida , is illustrated in Fig. 4.

Fig. 3. Amblyeleotris neumanni n. sp., female, Lolobau Island, New Britain. Photo by J. E. Randall.

Fig. 4. Amblyeleotris neumanni n. sp . female, Lolobau Island, New Britain. Photo by J. E. Randall.

23 aqua vol. 11 no. 1 - 2006 Amblyeleotris neumanni , a new species of shrimp goby from New Britain

Amblyeleotris neumanni is most similar to A. masuii Aonuma and Yoshino, described from Okinawa in 1996. It was illustrated in colour by Hayashi and Shi - ratori (2003: 139, lower fig.) and Senou et al . (2004: 328). The two species share the same fin ray counts, nearly the same high number of scales in longitudinal series, and most morphometric data. Both have a lanceolate caudal fin and the pelvic fins fully joined medially with a well developed frenum. There is also much similarity in life colour. Amblyeleotris masuii differs in having a shorter snout, 16.4-18.6% HL, compared to 23.0-23.5% in A. neumanni , with a steeper dorsal snout profile (about 60°, compared to about 40° in neumanni ). The male of masuii lacks the filamentous third to fifth dorsal spines that are found on neumanni . Also, masuii is evidently a larger species, the type specimens mea - suring 61.0-71.7 mm SL, compared to the largest neumanni , 54.9 mm. The salient colour differences are the orange and blue spots and orange-tipped rays of the dorsal fins of neumanni (no obvious spots on these fins in masuii , the first dorsal edged in dark red - dish brown, and the second dorsal rays tipped in dark reddish brown), the dark brown band behind the mid - dle of the eye of neumanni , and the four different coloured bands on the base of the anal fin of neu - manni (only a single orange line in masuii).

Acknowledgements We thank Mike Neumann for his assistance in col - lecting and photographing the type specimens of the new species, Arthur Anker for the identifications of the symbiotic alpheid shrimps, and Gerald R. Allen, David W. Greenfield and Helen A. Randall for their review of the manuscript.

References Aonuma, Y. & T. Yoshino. 1996. Two new species of the genus Amblyeleotris (Pisces: Gobiidae) from the Ryukyu Islands, Japan. Ichthyological Research, 43 (2): 161-168. Bleeker , P. 1874. Notice sur les genre Amblyeleotris, Valenciennesia et Brachyeleotris, Verslagen en Mededeelingen der Koninklijke Akademie van Wetenschappen , 8: 372-376. Hayashi, M. & T. Shirator i, 2003. Gobies of Japanese Waters . 223 pp. TBS Buritanica, Tokyo (in Japanese). Masuda, H., Amaoka, K., Araga, C., Uyeno, T. & T. Yoshino (Eds.) . 1984. The Fishes of the Japanese Archipelago . Vol. 1 (text xxii + 437 pp) and vol. 2 (plates). Tokai University Press, Tokyo. Randall, J. E . 2005. Five new shrimp gobies of the genus Amblyeleotris from islands of Oceania. aqua, Journal of Ichthyology and Aquatic Biology, 8 (2): 61-78. Senou, H., Suzuki, T., Shibukawa, K. & K. Yano . 2004 . A Photographic Guide to the Gobioid Fishes of Japan. 534 pp. Heibonsha, Ltd., Tokyo (in Japan ese). aqua vol. 11 no. 1 - 2006 24 aqua, Journal of Ichthyology and Aquatic Biology

Descriptive anatomy of Iso rhothophilus (Ogilby), with a phylogenetic analysis of Iso and a redefinition of Isonidae (Atheriniformes)

Basim Saeed, Walter Ivantsoff and Aarn

Departmentl of Biological Sciences, Macquarie University 2109, Australia. E-mail: [email protected]

Accepted: 03.06.2005

Keywords niformes) si dimostra distinta dai Notocheiridae Isonidae, phylogenetic analysis, anatomy, osteology Schultz, 1950.

Abstract Introduction The musculoskeletal anatomy of Iso rhothophilus is Isonids are small fishes that have been infrequently described. A phylogenetic analysis of the genus, using collected in surf, usually near rocks, at the margins of eleven anatomical characters, indicates that the sys - the Pacific, Indian and Atlantic Oceans. The history of tematic hierarchy is (Iso flosmaris (I. nesiotes (I. the genus Iso and the designation of the family rhothophilus (I. hawaiiensis, I. natalensis)))). Isonidae Isonidae are fraught with confusion. Ogilby (1895) Rosen, 1964 is redefined and, on the basis of twenty described the first species of the “flower of the wave” characters (autapomorphic within Atheriniformes) (as it was subsequently commonly called) as Tropi - shown to be distinct from Notocheiridae Schultz, 1950. dostethus rhothopilus from Maroubra Bay, NSW, Australia. In 1901, Jordan and Starks named another Zusammenfassung species as Iso flosmaris from Japan. A few years later Beschrieben wird die Anatomie des Muskelskeletts Waite (1904) indicated that Iso was indistinct from von Iso rhothophilus. Nach einer phylogenetischen Tropidostethus, then, in 1919, Jordan and Hubbs Analyse der Gattung anhand von elf anatomischen pointed out that Tropidostethus was preoccupied by a Merkmalen muss man auf die folgende systematische genus of insects thus determining the validity of the Hierarchie schließen: (Iso flosmaris (I. nesiotes (I. name Iso. Other species ascribed to the genus were rhothophilus (I. hawaiiensis, I. natalensis)))). Die I. natalensis Regan, 1919, I. flosindicus Herre, 1944, Isonidae Rosen, 1964, werden neu definiert und ihre I. hawaiiensis Gosline (1952) and I. nesiotes Saeed, Unterscheidung zu den Notocheiridae Schultz, 1950, Ivantsoff & Crowley, 1993 . A description of Noto- auf der Grundlage von 20 Merkmalen (autapomor - cheirus hubbsi by Clark in 1937 was noted by Hubbs phisch innerhalb der Atheriniformes) festgestellt. (1944) as a species which was Iso- like but according to him was likely to be an offshoot of Atherinopsinae. Résumé Further confusion arose with the description of a On décrit l'anatomie musculo-squelettique d' Iso subfamily of Tropidostethinae by Schultz in 1948. rhothophilus. Une analyse phylogénétique du genre, According to him this subfamily included Notocheirus, à l'aide de onze caractéristiques anatomiques, mon - Tropidostethus and Iso, presumably considering that tre que la hiérarchie systématique est (Iso flosmaris the type genus of the subfamily was Tropidostethus. (I. nesiotes (I. rhothophilus (I. hawaiiensis, I. natalen - In 1950 Schultz appears to have realised that Tropi - sis)))). Isonidae Rosen, 1964 est redéfini et, à l'aide dostethus was preoccupied, having substituted the de vingt caractéristiques (autapomorphiques pour les name with Tropidosthetops, at the same time chang - Atheriniformes), on montre sa différence par rapport à ing the subfamilial name to Notocheirinae, and stating Notocheiridae Schultz, 1950. that the new subfamily was based on the genus Notocheirus Clark. Sommario In a review of Iso Jordan and Starks (Said, 1983), L’anatomia muscoloscheletrica di Iso rhothophilus è five species were considered valid, and the genus descritta in dettaglio. Un’analisi filogenetica del ge- was considered distinct from Notocheirus Clark. Said nere, eseguita con undici caratteri anatomici, indica (1983) noted that Isonidae Rosen, 1964 was erected che la gerarchia sistematica è (Iso flosmaris (I. nesio - on the basis of examination of Notocheirus hubbsi tes (I. rhothophilus (I. hawaiiensis, I. natalensis)))). La Clark, a monotypic genus previously assigned type famiglia Isonidae Rosen, 1964 viene ridefinita e, sulla status for Notocheirinae Schultz, 1950. base di venti caratteri (autapomorfici entro gli Ateri - In a review of the systematic position of Isonidae

25 aqua vol. 11 no. 1 - 2006 Descriptive anatomy of Iso rhothophilus (Ogilby), with a phylogenetic analysis of Iso and a redefinition of Isonidae (Atheriniformes)

(Saeed et al ., 1993), the family was considered to be myodomes present; mesethmoid morphology unique, distinct from Notocheiridae. However, the most recent nasal septum formed by ethmoid and mesethmoid; review of Atheriniformes (Dyer and Chernoff, 1996) parietals not contributing to posttemporal fossa; ante - contested that view. The present authors therefore rior vertebral parapophyses with lateral ridges; inter - decided to further examine isonid diagnostics and dorsals absent; cleithrum dorsal enclosure absent; relationships. membrane just posterior to genital opening; and pelvic posteromedial process elongate. Materials Type material examined for each species is listed Iso Jordan and Starks, 1901 first, followed by materials prepared for osteological Tropidostethus Ogilby, 1895:323, type species: Tropi - and/or micrographic examination. The number and dostethus rhothopilus by monotypy, preoccupied by size of specimens follows institutional identification. Tropidostethus Philippi, 1863 in Orthoptera. Waite, Institutional abbreviations are as follows: AMS: Aus - 1904a:234; Schultz, 1948:26; Munro, 1958: 99. tralian Museum, Sydney; BMNH: British Museum Iso Jordan and Starks, 1901: 205, type species: Iso flos - (Natural History); BPBM: CAS: Californian Academy maris by monotypy. Waite, 1904b:21; Jordan and of Science; MU: Macquarie University, Sydney; Hubbs, 1919: 47; Regan, 1919: 200; McCulloch, 1929: SOSC: Smithsonian Oceanographic Sorting Center, 110; Herre, 1944: 47; Gosline, 1952: 49; Golvan , Washington DC; USNM: Smithsonian Institution, 1959: 73; Rosen, 1964: 227; Smith, 1965: 603. Washington, DC; ZUMT: Department of Zoology, Uni - Tropidostethops Schultz, 1950: 150, type species: versity of Tokyo. Tropidostethus rhothophilus Ogilby, 1895 (replace - Notocheirus hubbsi : CAS 5525 (holotype), CAS ment name for Tropidostethus, which is preoccu - 5526 (paratype, disarticulated); MU I-307 (2, 32 pied). mmSL). Iso flosmaris : CAS SV6527 (holotype); CAS 7142 (10, paratypes); SOSC VGS-68-30 (4, 22-60 Iso rhothophilus (Ogilby, 1895) mm SL); ZUMT 5918 (1, 54 mm SL). Iso hawaiiensis : USNM 152759 (holotype); BPBM 15249 (2, External morphology paratypes); BPBM 10012 (1, 28 mm SL). Iso natalen - Body highly compressed, deepest at level of vertical sis : BMNH 1919.4.1.13 (holotype); CAS 3384 (3, 25- plane through origin of pectorals, tapering towards 35 mm SL); MU I-227 (3, 28-45 mm SL). Iso nesiotes : caudal peduncle (Fig.1). Ventral edge of abdomen AMS I241183-001 (holotype); BPBM 16718 (1, 22 tapering to keel-like form. Head small, truncate pos - mm SL); MU I-210 (5, 20-27 mm SL). Iso teriorly, snout round, mouth oblique. Premaxillary rhothophilus : BMNH 1897.3.20-22 (holotype); MU I- non-protractile, attached to head by mid-dorsal 122 (19, 21-55 mm SL); SOSC VGS-68-22 (2, 21-48 frenum. Maxillary slightly concave anteriorly. Lower mm SL); ZUMT 4158 (5, 38-46 mm SL). jaw deeply elevated posteriorly, placed medial to upper elements. Methods Premaxillary and dentary teeth small, well devel - Specimen clearing and staining, examination and oped, curving backwards into mouth. Vomerine teeth illustration techniques, and preparation for electron present in some specimens. Palatine teeth absent. micrography, follow Saeed et al. , 1993. Anatomical Gills opening widely. Gill rakers on first gill arch well terminology is based on Winterbottom (1974). developed. Anus just anterior to origin of anal fin. Phylogenetic analysis was based on characters Small membrane posterior to genital opening (Fig. 2). identified by comparison of specimens of five Iso spp. First dorsal fin originating approximately at vertical Notocheirus hubbsi was selected as the outgroup. through midbody, of III-VI small weak spines (Fig. 1). Character states were coded 0, 1 or 2 by the outgroup Second dorsal originating posterior to vertical through comparison method. All characters were considered anal origin, comprising spine, unbranched ray and 9- unordered, and all characters assigned equal weight. 16 branched rays. Anal fin comprising spine, The phylogeny was derived using a closest pair com - unbranched ray and 19-27 branched rays. Second parison method i.e. the two species with most shared dorsal and anal high anteriorly, tapering posteriorly. plesiomorphies were joined, this pair was then joined Pectoral fin comprising spine, unbranched ray and 11- to the next most similar species and so on. 14 branched rays. Pectoral compressed, broad, inserted high on body with dorsalmost ray originating Isonidae Rosen, 1964 above posteriormost point of opercle. Pectoral poste - Type genus Iso Jordan and Starks, 1901 (replacing rior angle rounded, covering origin of midlateral band. Notocheirus Clark, 1937). Pelvics compressed, inserted low on body. Scales in midlateral series 42-55. Body silver, dark Diagnosis midlateral band from axilla to tail, discontinuous on A monogeneric atheriniform family with the paras - caudal peduncle. Scales ovoid, absent from cranial phenoid concave rather than convex, parasphenoid region. Larger specimens to 60 mm SL. aqua vol. 11 no. 1 - 2006 26 Basim Saeed, Walter Ivantsoff and Aarn

Fig. 1. Iso rothophilus , MU I-122, 54 mm SL.

Fig. 2. Iso rothophilus , AMS 17766, 47.4 mm SL. Anal region in lateral aspect. Abbreviations: af: anal fin; an: anus; ge: genital opening; gm: genital membrane.

27 aqua vol. 11 no. 1 - 2006 Descriptive anatomy of Iso rhothophilus (Ogilby), with a phylogenetic analysis of Iso and a redefinition of Isonidae (Atheriniformes)

Musculoskeletal anatomy lage, ventral to condyle for lacrimal. Lateral ethmoid Cranium: Rostrum elongate (Fig. 3). Ethmoid carti - closely contacting contralateral structure, not forming lage partially exposed laterally to form facet for articu - articulation with parasphenoid. Lateral ethmoid medial lation with palatine. Lateral ethmoid with cartilaginous border with notch for olfactory nerve. condyle for articulation with lacrimal, supported dorsally Rostral cartilage bound to premaxilla dorsal process. and ventrally by osseous laminae. Lateral ethmoid with Dorsal mesethmoid absent. Vomer with bilateral max - cartilaginous process contacting palatoquadrate carti - illary condyles (Fig. 4). Parasphenoid enlarged poste -

A

B

Fig. 3. Iso rothophilus , MU I-122, 50.1 mm SL. Neurocranium in A: lateral aspect; B: ventral aspect. Abbreviations: bo: basioccipital; bs: basisphenoid; cp: palatine condyle; ec: ethmoid cartilage; eo: exoccipital; eoc: exoccipital crest; ep: epiotic; fm: foramen magnum; fr: frontal; gf: glossopharyngeal foramen; hf: hyomandibular facet; jc: jugular canal; lb: laminar portion of mesethmoid; lc: lacrimal; le: lateral ethmoid; mc: maxillary condyle; me: mesethmoid; mo: myodome opening; na: nasal; ob: otic bulla; oc: occipital condyle; of: olfactory nerve foramen; pa: parasphenoid; pc: epiotic crest; pe: pterotic; pf: palatine facet; pi: parietal; pj: pterotic projection on exoccipital; po: postorbital process; pr: prootic; ps: pterosphenoid; pt: posttemporal fossa; sc: supraorbital canal; so: supraoccipital; sp: sphenotic; st: subtemporal fossa; tc: temporal canal; vf: vagus foramen; vo: vomer. Scale bar: 1mm. aqua vol. 11 no. 1 - 2006 28 Basim Saeed, Walter Ivantsoff and Aarn

Fig. 4. Iso rothophilus , MU I-122, 50.2 mm SL. Scanning electron micrograph of neurocranium in ventral aspect. A: rostral portion; B: central portion. Abbreviations: ec: ethmoid cartilage; fr: frontal; lc: lacrimal condyle; le: lateral ethmoid; mc: maxillary condyle; pa: parasphe - noid; pf: palatine facet; pr: prootic; ps: pterosphenoid; vo: vomer. riorly, embedded in basioccipital. Parasphenoid ante - rior margin planar, interposed between ethmoid carti - lage and vomer. Parasphenoid ventral ridge com - pressed, blade-like, extending posteriorly. A Nasal irregular to triangular, separated from con - tralateral structure. Nasal cavity extending medially, separated from contralateral structure by thin ventro - median lamina of mesethmoid. Accessory nasal sac located medial to anterior infraorbitals. Frontals forming major portion of roof of skull. Frontal truncate anteriorly, not extending forward over nasal cavity, thereby partially exposing ethmoid. Frontal contacting mesethmoid but not lateral eth - moid. Interorbital portion of frontal separated from supraorbital laminae by deep sulcus forming supraor - bital sensory canal (Fig. 5). Parietal large. Supraoccipital drawn into long median process deep to frontals. Epiotic large, bearing crest for attachment of epaxial musculature (lesser crests also occurring on supraoccipital, exoccipital and pterotic). Posttem - poral fossa formed by pterotic and sphenotic, and in B part by epiotic and parietal, bounded laterally by osseous temporal canal. Temporal canal fused to pterotic, extending anteriorly to form junctions with Fig. 5. Iso rhothophilus , MU I-122, 50.1 mm SL. Neuro - supraorbital and infraorbital canals. Foramen situated cranium in A: dorsal aspect; B: caudal aspect. Abbrevi - at anterior temporal canal, superficial to sphenotic, ations: bo: basioccipital; ec: ethmoid cartilage; forming nexus between dilator operculi fossa and eo: exoccipital; eoc: exoccipital crest; ep: epiotic; canal. Dilator operculi fossa giving origin, in part, to fm: foramen magnum; fr: frontal; hf: hyomandibular face; lacrimal; lateral ethmoid; mesethmoid; levator operculi muscle. Anterior wall of fossa formed lc: le: me: mo: myodome opening; na: nasal; oc: occipital condyle; by sphenotic postorbital process. of: olfactory nerve foramen; pa: parasphenoid; pc: epi - Sphenotic and pterotic forming facets articulating otic crest; pe: pterotic; pf: palatine facet; pi: parietal; with hyomandibula. Subtemporal fossa medial and pt: posttemporal fossa; sc: supraorbital canal; deep to posttemporal fossa, giving origin to branchial so: supraoccipital; sp: sphenotic; tc: temporal canal; levator musculature. vf: vagus foramen; vo: vomer. Scale bar: 1mm.

29 aqua vol. 11 no. 1 - 2006 Descriptive anatomy of Iso rhothophilus (Ogilby), with a phylogenetic analysis of Iso and a redefinition of Isonidae (Atheriniformes)

AB

Fig. 6. Iso rhothophilus , MU I-122, 52 mm SL. A: skull and pectoral girdle in lateral aspect; B: superficial musculature and ligaments of the head. Abbreviations: a1, a2, aw: divisions of adductor mandibulae muscle; am: articularmaxillary ligament; ar: articular; br: branchiostegal rays; cl: cleithrum; co: coracoid; de: dentary; ds: dermosphenoid; df: dilator fossa; dop: dilator operculi muscle; dpa: epaxial musculture; fr: frontal; hp: hypopalatine arch; hy: hyomandibula; i1-3: infraorbitals 1-3; io: interopercle; la: labial ligament; lp: levator arcus palatini muscle; lop: levator operculi muscle; lt: lacrimal tendon; ma: maxilla; me: mesethmoid; ms: mesopterygoid; mp: metapterygoid; mt: maxillary tendon; na: nasal; op: opercle; phy: protractor hyoideus muscle; pi: parietal; pm: premaxilla; po: posttemporal; pr: preopercle; pt: posttemporal fossa; pq: palatoquadrate cartilage; ra: proximal radials; re: retroarticular; ro: rostral cartilage; sc: scapular; sf: scapular foramen; shy: sternohyoideus ; so: subopercle; sp: sphenotic. Scale bar: 1 mm.

Pterosphenoid separated from contralateral struc - nal processes formed about premaxilla dorsal ture, lateral margin contacting frontal. Pterosphenoid, process. Maxilla joined anteriorly to contralateral and ventral ridge originating on frontal, supporting structure ligamentously. Premaxilla process providing membrane enclosing anterior lobe of brain. Mem - attachment for articularmaxillary ligament (Fig. 7), brane attached posteriorly to prootic and basisphe - originating from articular lateral face. Rostral cartilage noid. Basisphenoid bearing bilateral posterolateral supporting premaxilla dorsal process. No ligaments rami contacting prootic. between maxilla and palatine. Orbits separated medially by septum attached to Ethmomaxillary ligament originating on mesethmoid, parasphenoid. Ventromedian projection from prootics inserting on maxilla dorsal process. Labial ligament extending between bilateral rami of basisphenoid. between anterolateral face of dentary and distal Prootic jugular canal deep to osseous trabeculum. angles of maxilla and premaxilla. Coronomaxillary lig - Posterior jugular opening confluent with emanation of ament absent. hyomandibular nerve. Adductor mandibulae muscle inserting in triangular Otic bulla small, extending ventrolaterally to paras - recess dorsal to Meckelian cartilage between dentary phenoid. Pterotic bearing projection superficial to and articular. Meckelian cartilage elongate (Fig 8), exoccipital. joining dentary and articular medially. Coronomecke - Occipito-vertebral articulation formed by spherical lian bone, on dorsal surface of Meckelian cartilage basioccipital condyle and paired small, separate adjacent to articular, acting as insertion for tendon of exoccipital condyles. adductor mandibulae muscle . Articular with retrorse Jaws: Premaxilla with moderate symphysis, alveolar condyle on posterodorsal facet serving as insertion for ramus slightly smaller than maxilla (Fig. 6). Premaxilla ligament from quadrate. Retroarticular bone small, at postmaxillary processes supporting skin fold between angle of lower jaw, overlapping medially with articular. premaxilla and maxilla. Premaxilla teeth forming sin - Suspensorium: Comprising palatoquadrate arch, gle row. Maxilla narrow, positioned lateral to premax - ectopterygoid, mesopterygoid, ectopterygoid, sym - illa. Maxilla proximal angle narrow, distal angle broad. plectic, hyomandibula and preopercle (Fig 9). Two Maxilla bearing large condyle for articulation with articulations formed with cranium, bilaterally: anterior vomer, biconcave submaxillary meniscus interposed articulation with palatine, posterior articulation with between maxilla and vomer. Maxilla dorsal and inter - hyomandibula. Palatine and quadrate joined about aqua vol. 11 no. 1 - 2006 30 Basim Saeed, Walter Ivantsoff and Aarn

Fig. 7. Iso rhothophilus , MU I-122, 50.8 mm SL. Upper jaw and nasal area (lacrimal removed). Abbreviations: em: eth - momaxillary ligament; lb: laminar portion of mesethmoid; lc: lacrimal condyle; le: lateral ethmoid; ma: maxilla; mi: mesethmoid; na: nasal; nm: nasomaxillary ligament; pa: palatine; pf: palatine facet; pm: premaxilla; pq: palatoquadrate cartilage; ro: rostral cartilage; tc: ethmoid cartilage. Scale bar: 1 mm. interposed cartilage, cartilage also contacting eth - moid. Palatine cartilage progressively ossifying. Pala - tine maxillary process comprising cylindrical bone with cartilaginous core. Palatine bearing posteroven - tral process overlapping ectopterygoid. Mesopterygoid expansive, thin, forming ventrome - dian limit of orbit. Metapterygoid and quadrate joined about narrow interposed cartilage. Quadrate and A symplectic with elongate articulation, both bones with broad ventral facets articulating with preopercle. Metapterygoid not articulating with cranium. Hyomandibula with ventral projection joined to pre - opercle ventral ramus. Hyomandibula articulating dor - sally with cranium, hyomandibula posterior condyle articulating with opercle. Opercular series: Comprising opercle, interopercle and subopercle. Opercle large, elongate, bearing gle - noid fossa articulating with hyomandibula. Dilator operculi muscle inserting on medial face of triangular dilator process of sphenotic and pterotic. Levator B operculi muscle inserting on dorsal and medial sur - faces of opercle. Interopercle wide, bearing dorsal process. Interopercle and branchiostegal rays covering Fig. 8. Iso rhothophilus , MU I-122, 52 mm SL. Scanning gills ventrally. Interopercle joined to lower jaw ligamen - electron micrograph of A: right premaxilla in lateral tously. Interopercle joined to ceratohyal by short liga - aspect; B: lower jaw in medial aspect. Abbreviations: ar: ment. Subopercle elongate, forming portion of gill cover. articular; dc: coronoid process; de: dentary; me: Neurosensory network: Sensory canals developed Meckel’s cartilage; re: retroarticular. Scale bar: 1 mm.

31 aqua vol. 11 no. 1 - 2006 Descriptive anatomy of Iso rhothophilus (Ogilby), with a phylogenetic analysis of Iso and a redefinition of Isonidae (Atheriniformes)

Fig. 9. Iso rhothophilus , MU I-122, 50.1 mm SL. Right suspensorium, lower jaw and opercular series in medial aspect. Abbreviations: ar: articular; cm: coronomeckelian; dc: coronoid process; de: dentary; ec: ectopterygoid; fm: foramen of hyomandibular nerve; hy: hyomandibula; io: interopercle; me: Meckel’s cartilage; ms: mesopterygoid; mt: metaptery - goid; on: notch in opercle; op: opercle; pa: palatine; pq: palatoquadrate cartilage; pr: preopercle; qa: quadrate; re: retroarticular; so: subopercle; sy: symplectic. Scale bar: 1 mm. on nasal, frontal, pterotic, posttemporal, preopercle, urohyal; and bilaterally paired dorsal and ventral infraorbital, dentary and articular bones, lateral line hypohyals, anterior and posterior ceratohyals, inter - absent. Infraorbital series comprising four elements. hyal, six branchiostegal rays, three hypobranchials, Lacrimal (infraorbital 1) horizontal axis elongate, five ceratobranchials, four epibranchials and three located superficial to premaxilla (when mouth closed), pharyngobranchials (Fig.10). bearing facets for articulation with lateral ethmoid, Basihyal rhomboidal, ossified posteriorly, strongly posterior border with recess contacting lateral eth - attached to hypohyals. Basihyal cartilaginous anterior moid lacrimal condyle (Fig. 3) . Lacrimal medial face portion supporting small tooth plate. Urohyal with bearing osseous shelf, coursing ventral to nasal sac. bilateral posterodorsal processes and median pos - Second infraorbital planar, irregular. Third infraorbital teroventral process. Three median basibranchials cylindrical. Dermosphenotic at posterolateral margin coursing antero-posteriorly from basihyal to irregular of orbit, contacting sphenotic postorbital process. cartilaginous nodule. Posteriormost two basibran- Supraorbital canal coursing antero-posteriorly from chials fused dorsally to respective tooth plates. nasal medial margin, along frontal sulcus, curving lat - Hyoid bar comprising dorsal and ventral hypohyals, erally to confluence with temporal and infraorbital and anterior and posterior ceratohyals. Hyoid bar canals. Temporal and preopercular canals not con - joined to contralateral structure by strong ligament joined. Mandibular sensory canal coursing along ven - from medial face of hypohyals, coursing ventral to tral border of articular and dentary. Canal openings on basihyal. Anterior and posterior ceratohyals joined dentary ventral surface. about dentate suture. Anterior ceratohyal ventral sur - Hyobranchial apparatus: Comprising median un- face giving origin to two anteriormost small bran - paired glossohyal, basihyal, three basibranchials and chiostegal rays, then two large rays rising from ante - aqua vol. 11 no. 1 - 2006 32 Basim Saeed, Walter Ivantsoff and Aarn

A

B

C

Fig. 10. Iso rhothophilus , MU I-122, 50.1 mm SL. A: Hyobranchial apparatus in dorsal aspect, right side elements folded out; B: Urohyal in dorsal aspect (upper element) and lateral aspect (lower element): C: Hyoid bar in lateral aspect. Abbre - viations: an: anterior ceratohyal; at: urohyal anterodorsal process; b1-3: basibranchials 1-3; bh: basihyal; br: branchioste - gal ray; bt: free branchiostegal toothplate; c1, c5: ceratobranchials 1 and 5; ch: ceratohyal; cn: cartilage nodule; cr: cra - nial condyle ; dh: dorsal hypohyal; e1-4: epibranhials 1-4; gr: gill rakers; h1-3 hypobranchials 1-3 : ih: interhyal; lt: process for hypohyal ligament; p1-3: pharyngobranchials 1-3; pd: urohyal posterodorsal process; po: posterior ceratohyal; pv: uro - hyal posteroventral process; un: epibranchial uncinate process; vh: ventral hypohyal. Scale bar: 1 mm.

33 aqua vol. 11 no. 1 - 2006 Descriptive anatomy of Iso rhothophilus (Ogilby), with a phylogenetic analysis of Iso and a redefinition of Isonidae (Atheriniformes)

A

B

C

Fig. 11. Iso rhothophilus , MU I-122, 50.1 mm SL. A: Anterior vertebrae; B: Last precaudal vertebra and anterior caudal vertebrae; C: Caudal skeleton. Abbreviations: ad: anterior dorsal zygapophysis; bp: parhypural basal process; ca: first caudal vertebra; ep: epural; er: epipleural; h1 + 2: fused hypurals 1 + 2; h3+4: fused hypurals 3+4; ha: haemal spine; me: median plate; ns: neural spine; pa: parapophysis; pd: dorsal postzygapophysis; ph: parhypural; pr: pleural rib; pu2: second preural vertebra; pv: posterior ventral zygapophysis; tc: terminal half-centrum; ul: uroneural; vz: anterior ventral zygapophysis. Scale bar: 1 mm. aqua vol. 11 no. 1 - 2006 34 Basim Saeed, Walter Ivantsoff and Aarn

Fig. 12. Iso rhothophilus , MU I-122, 49 mm SL. Pelvic girdle in A: lateral aspect; B: detail. Abbreviations: al: pelvic pos - teromedial process; ap: adductor profundus pelvicus muscle; as: abductor superficialis pelvicus muscle; pb: pelvic bone; pv: posteroventral process. Scale bar: 1 mm.

35 aqua vol. 11 no. 1 - 2006 Descriptive anatomy of Iso rhothophilus (Ogilby), with a phylogenetic analysis of Iso and a redefinition of Isonidae (Atheriniformes)

Fig. 13. Notocheirus hubbsi , MU I-307.

Fig. 14. Notocheirus hubbsi , CAS 5526 (paratype) premaxilla. Scale bar: 1 mm.

Fig. 15. Iso rhothophilus , MU I-122, 49 mm SL. Premaxilla. Fig. 16. Iso natalensis , MU I-227, 40.1 mm SL. Premaxilla. Abbreviation: ps: postmaxillary process. Scale bar: 1 mm. Abbreviation: ps: postmaxillary process. Scale bar: 1 mm. aqua vol. 11 no. 1 - 2006 36 Basim Saeed, Walter Ivantsoff and Aarn

Fig. 18. Iso hawaiiensis , BPBM 10012, 28 mm SL. Scan - Fig. 17. Iso natalensis , MU I-227, 45.1 mm SL. Scanning ning electron micrograph of neurocranium in ventral electron micrograph of premaxilla. aspect. Abbreviations: ap: pterosphenoid anterior process; fr: frontal; hf: hyomandibular facet; mp: pterosphenoid medial process; pa: parasphenoid; pr: prootic; ps: pterosphenoid; sp: sphenotic.

Fig. 19. Iso rhothophilus , SOSC VGS-68-22, 48.3 mm SL. Scanning electron micrograph of pterosphenoid. Abbrevi - ations: ap: pterosphenoid anterior process; fr: frontal; mp: pterosphenoid medial process; ps: pterosphenoid; sp: sphenotic. rior ceratohyal lateral surface. Two posteriormost rays rising from posterior ceratohyal lateral surface. Elongate interhyal bone forming in interhyal ligament between posterior ceratohyal and hyomandibula. Posterior ceratohyal bearing lateral facet giving rise to ligament to interopercle. Anterior three gill arches each of hypobranchial, cer - Fig. 20. Notocheirus hubbsi , MU I-307, 32.3 mm SL. Cra - atobranchial, epibranchial and pharyngobranchial, nium in dorsal aspect. Abbreviations: ec: ethmoid carti - fourth arch of ceratobranchial and epibranchial, fifth lage; ep: epiotic; fr: frontal; le: lateral ethmoid; pi: parietal; arch comprising single ceratobranchial. Gill rakers of po: postorbital process; so: supraoccipital; sp: sphenotic; first gill arch elongate. Ceratobranchials elongate, tc: temporal canal; vo: vomer.

37 aqua vol. 11 no. 1 - 2006 Descriptive anatomy of Iso rhothophilus (Ogilby), with a phylogenetic analysis of Iso and a redefinition of Isonidae (Atheriniformes) arcuate, ventral surface bearing process for attach - form haemal arches of caudal vertebrae. First haemal ment of pharyngeal musculature. Fifth cerato - arch emanating from first caudal vertebra, anterior branchial posterolateral angle produced as site of parapophysis directed ventromedially to fuse with con - attachment for posterior levator muscle. tralateral structure, produced as haemal spine. Hypobranchials fused dorsally to respective tooth Total pleural ribs 12-14, first rib contacting third ver - plates. Basibranchial tooth plate dorsal to, but not tebra. Epural articulating with respective vertebral fused with, third hypobranchial. Tooth plates adjacent body posterodorsal to base of parapophysis. First epi - to, but not fused with, first four ceratobranchials and pleural contacting dorsolateral aspect of parapoph - all epibranchials. Fifth ceratobranchial fused to large ysis of third vertebra, coursing posterodorsally to ribs. lower pharyngeal toothplate. Intervertebral articulations mediated by zygapophy - Axial skeleton: Total vertebrae 38-45, precaudal 14- ses. Dorsal zygapophyses prominent on anterior ver - 18, caudal 22-28. First vertebra with reduced neural tebrae, ventral zygapophyses present on posterior spine (Fig. 11), articulating with cranium. Neural arch precaudal and caudal vertebrae. rising from anterodorsal portion of respective vertebral Penultimate vertebra bearing very large haemal centrum. Neural arch of vertebra 2-6/7 developed as spine. Terminal half-centrum supporting hypurals and expanded neural plate. Precaudal vertebrae with para - parhypural. Parhypural fused with hypurals 1 and 2, pophyses rising from anteroventral portion of centrum. separated by horizontal gap from fused hypurals 3 Anterior parapophyses small, directed laterally, antero - and 4. Uroneural coursing posterodorsally from termi - posterior series becoming larger, directed posteriorly to nal half-centrum, epural anterior to uroneural.

Fig. 21. Iso flosmaris , SOSC VGS-68-30, 52 mm SL. Fig. 22. Iso hawaiiensis , BPBM 10012, 28 mm SL. Scanning electron micrograph of posterior neurocranium Scanning electron micrograph of portion of neurocra - in dorsal aspect. Abbreviations: ep: epiotic; fr: frontal; nium. Abbreviations: eo: exoccipital; ic: intercalar; pc: epiotic crest; pi: parietal; so: supraoccipital. pe: pterotic.

Fig. 23. Notocheirus hubbsi , CAS 5526 (paratype). Urohyal. Abbreviations: at: anterodorsal process; lt: process for hypohyal ligament; pd: posterodorsal process. Scale bar: 1 mm. aqua vol. 11 no. 1 - 2006 38 Basim Saeed, Walter Ivantsoff and Aarn

Table I. Data matrix showing distribution of seven character states across outgroup and five ingroup taxa.

Notocheirus I. flosmaris I. nesiotes I. rhothophilus I. hawaiiensis I. natalensis

1 002111 2 000102 3 001111 4 000001 5 011101 6 010011 7 000010 8 001111 9 001000 10 000010 11 010111

Table II. The number of shared plesiomorphic character states between pairs of six taxa.

I. flosmaris I. nesiotes I. rhothophilus I. hawaiiensis I. natalensis Notocheirus 865 4 3 I. flosmaris 44 3 3 I. nesiotes 422 I. rhothophilus 23 I. hawaiiensis 1

Fig. 24. Iso hawaiiensis , BPBM 10012, 28 mm SL. Anterior vertebrae. Abbreviations: ad: anterior dorsal zygapophysis; er: epipleural; me: median plate; ns: neural spine; pa: parapophysis; pr: pleural rib. Scale bar: 1 mm.

39 aqua vol. 11 no. 1 - 2006 Descriptive anatomy of Iso rhothophilus (Ogilby), with a phylogenetic analysis of Iso and a redefinition of Isonidae (Atheriniformes)

Fins and girdles: Each spine of first dorsal supported thrum in most specimens. Supracleithrum absent. by individual basal radial comprising fused proximal Posttemporal trabecular, arcuate, superficial to epiotic and medial elements. No radials between dorsal fins. dorsal crest. Dorsal postcleithrum absent, ventral Each element of second dorsal and supported by postcleithrum contacting coracoid. basal radial and non-ossified distal radial, basal radi - Pelvic fin without distal radials, fin elements sup - als also supporting preceding element. ported directly (Fig. 12). Adductor profundus pelvicus Pectoral fin with four proximal radials supporting fin muscle on dorsomedial aspect of pelvic, abductor rays, dorsal radial fused to scapula (Fig. 6A). Scapu - superficialis on lateral aspect. Pelvic posteromedial lar foramen between scapula and cleithrum. Coracoid process elongate, extending dorsally. Pelvic pos - contacting cleithrum and scapula dorsally, ventrally teroventral process fused to contralateral structure. forming medial shelf and additional contact with clei - Character analysis The following characters, which differed between the outgroup and one or more ingroup taxa, were identified: 1. Midlateral band. In Notocheirus (Fig. 13) and I. flosmaris , the midlateral band is continuous. In I. hawaiiensis , I. natalensis and I. rhothophilus (Fig. 1), the band is discontinuous on the caudal peduncle, resulting in the presence of a ‘spot’ near the tail. In I. nesiotes (Saeed et al ., 1993; Fig. 1), the band is absent from the posterior part of the caudal peduncle. (0 = midlateral band continuous; 1 = band discontin - uous; 2 = band foreshortened). 2. Premaxilla symphyseal facet. The premaxillae have large symphyseal portions in Notocheirus (Fig. 14), I. flosmaris , I. hawaiiensis and I. nesiotes. The symphysis is reduced in I. rhothophilus (Fig. 15) and small in I. natalensis (Fig. 16). (0 = premaxilla symphyseal facet large; 1 = symph - ysis reduced ; 2 = symphysis small). 3. Premaxilla posterior angle expansive. In Notocheirus (Fig. 13) and I. flosmaris , the posterior angle of the premaxilla is broad. In other taxa (Figs. 15, 16) it is narrow/pointed. (0 = premaxilla posterior angle broad; 1 = posterior angle narrow). 4. Premaxilla bearing laterally-placed teeth. There are no teeth lateral to the premaxilla in all taxa except I. natalensis (Figs. 16, 17). (0 = Premaxilla lateral teeth present; 1 = teeth exter - nal to premaxilla). 5. Pterosphenoid anteromedial facet. In Notocheirus and I. hawaiiensis (Fig. 18) the pterosphenoid has a stright anteromedial facet. In other taxa this facet is curved (Fig. 19). (0 = pterosphenoid anteromedial facet linear; 1= facet lunate). 6. Parietal. The parietal is large in Notocheirus (Fig. 20), I. nesiotes and I. rhothophilus (Fig. 5), and small in the other taxa (Fig. 21). (0= parietal large; 1= parietal small). 7. Intercalar. The intercalar is absent in all taxa except I. hawaiiensis (Fig. 22). (0 = intercalar absent; 1= intercalar present). Fig. 25. Notocheirus hubbsi , CAS 5526 (paratype). Pec - 8. Basihyal teeth. There are no teeth on the basihyal toral girdle. Abbreviations: cl: cleithrum; co: coracoid; of Notocheirus and I. flosmaris . Other taxa have teeth pc: postcleithrum; po: posttemporal; ra: proximal radial; on the basihyal (Fig. 10). sc: scapular; sf: scapular foramen. Scale bar: 1mm. (0 = basihyal teeth absent; 1 = teeth present). aqua vol. 11 no. 1 - 2006 40 Basim Saeed, Walter Ivantsoff and Aarn

Table III. Osteological differences between Iso and Notocheirus . Autapomorphic states (within Atheriniformes) indicated A.

Iso Notocheirus 1 Scales elongate, denticulate (Said, 1983). A 2 Premaxilla symphysis bullous, premaxillary teeth distribution unique (Rosen, 1964). A 3 Mesethmoid absent, nasal cavities separated by ethmoid (Said, 1983). A 4 Mesethmoid morphology unique, nasal septum formed by ethmoid and mesethmoid (Saeed et al. , 1994). A 5 Vomer absent (Rosen, 1964). A 6 Parasphenoid ventromedian process (Rosen, 1964). A 7 Parasphenoid concave rather than convex, parasphenoid myodomes present (Said, 1983). A 8 Parietals present but not contributing to posttemporal fossa (Saeed et al. , 1994). A 9 Palatine reduced (Rosen, 1964). A 10 First epibranchial absent (Said, 1983). A 11 Anterior vertebral parapophyses with lateral ridges (Said, 1983). A 12 Interdorsals absent (Dyer and Chernoff, 1996). A 13 Epural absent (Rosen, 1964). A 14 Posterior pterygiophores of second dorsal and anal fins on individual unshared cartilage support (Dyer and Chernoff, 1996). A 15 Scapula and coracoid dorsal to midline (Rosen, 1964), pectoral inserted dorsally (Saeed et al. , 1994). A 16 Cleithrum dorsal enclosure absent (Dyer and Chernoff, 1996). A 17 Pectoral spur absent (Dyer and Chernoff, 1996). A 18 Membrane posterior to genital opening (Said, 1983). A 19 Pelvic posteromedial process elongate (Saeed et al. , 1994). A 20 Pelvic posteroventral process absent (Said, 1983), pelvic fin morphology unique (Saeed et al. , 1994) A

9. Urohyal posterodorsal process. In all taxa the uro - Discussion hyal posterodorsal process is directed posterodorsally The hierarchy of relationships in Iso was I. flosmaris (Figs. 10B, 23) except in I. nesiotes , in which the (I. nesiotes (I. rhothophilus (I. hawaiiensis, I. natalen - process is directed dorsally (Saeed et al. , 1993; Figs. sis))). This scheme postulates 5 reversals, while pro - 2Ca, 2Cb). viding deep analysis of the genus. Said (1983) exam - (0 = urohyal posterior process directed posterodor - ined five species (and a number of putative hybrid sally; 1 = process directed dorsally). forms), and used cluster and discriminant function 10. Anterior vertebral dorsal postzygapophyses. All analyses of osteological and/or morphological and/or taxa have well-defined processes (Fig. 11) except I. meristic data, to examine relationships within Iso . hawaiiensis (Fig. 24) in which the processes are not Although no two hierarchies were identical, the major - discernable. ity of schemes supported (I. flosmaris ((I. rho- (0 = anterior vertebral dorsal postzygapophyses well thophilus, I. natalensis), (I. hawaiiensis, I. nesiotes))). defined; 1 = processes not defined). The present scheme, based on a phylogenetic 11. Coracoid. The coracoid is relatively small in approach, supports the position of I. flosmaris as a Notocheirus (Fig. 25) and I. nesiotes (Saeed et al. , sister group to the other four species, but is otherwise 1993; Figs. 2Da, 2Db), and large in other taxa. dissimilar. (0 = coracoid small; 1 = coracoid large). Isonidae was erected by Rosen (1964) with eleven Eleven characters thus identified were entered into a osteological characters. Said (1983) described 33 data matrix (Table I). A second matrix was then devel - morphological and/or anatomical characters differing oped to show the number of shared plesiomorphies between Iso and Notocheirus . Subsequently Saeed et between pairs of taxa (Table II). This was used to al. (1994) erected Atherinopsoidea, comprising generate the cladogram (Fig. 26). (Atherinopsidae, Isonidae, Notocheiridae), with five

41 aqua vol. 11 no. 1 - 2006 Descriptive anatomy of Iso rhothophilus (Ogilby), with a phylogenetic analysis of Iso and a redefinition of Isonidae (Atheriniformes)

Fig. 26. Phylogenetic relationships of Iso . Node A. (Isonidae/Notocheiridae). Discussed by Saeed et al . (1994), Dyer and Chernoff (1996). Node B. (Isonidae). See Table III, Characters 5, 6, 11. Node C. Characters 1(2), 3, 6(R), 8, 9, 11(R). Node D. Characters 1(1), 2(1), 9(R), 11. Node E. Character 6. Node F. Iso hawaiiensis . Characters 2(R), 5(R), 10. Node G. Iso natalensis . Characters 2(2), 4. osteological characters, distinguishing this group from between Iso and Notocheirus commenced before the Atherinoidea. Notocheiridae was diagnosed with land masses of present day Africa and the Americas seven characters, while Isonidae was diagnosed with were conjoined. It is to be hoped that comparisons of six characters. molecular sequences will advance the present knowl - Dyer and Chernoff (1996) considered Iso and edge of isonid systematics. Notocheirus to be more derived than Atherinopsidae, and a sister group to the so-called infraorder Ather - Acknowledgements ines (comprising four families Melanotaeniidae, Athe - We wish to thank Ms Betty Thorn and Mr. Ron Old - rionidae, Phallostethidae and Atherinidae). On the field (Macquarie University) for preparation of some basis of examination of I. natalensis , I. rhothophilus illustrations and micrographs, and staff of institutions and N. hubbsi , and consideration of previous studies, who kindly facilitated loans of specimens. (Iso , Notocheirus ) Notocheiridae was diagnosed with three autapomorphies: supracleithrum absent; pelvic References dorsolateral process elongate, attaching to pleural rib Clark, H. W. (1937). New fishes from Templeton by elongate ligament; and body shape unique. Iso Crocker Expedition of 1934-35. Copeia, 2: 88-91. was diagnosed with six autapomorphies and four Dyer, B. S. & B. Chernoff . (1996). Phylogenetic rela - additional characters, Notocheirus was diagnosed tionships among the atheriniform fishes (Teleostei, with seven autapomorphies and six additional charac - Atherinomorpha). Zoological Journal of the Lin - ters. Thus, Dyer and Chernoff (1996) identified 23 naean Society, London, 117: 1-69. anatomical differences between Iso and Notocheirus . Golvan, Y. J. 1959. Catalogue systématique des Table III lists twenty autapomorphies of either Iso or noms de genres de poissons actuels, de la 10 e édi - Notocheirus the present authors have been able to tion “Systema naturae” de Charles Linné jusqu’à la verify, following Rosen (1964), Said (1983), Saeed et fin de l’année 1959 . Masson et Cie, Paris 227pp. al . (1994) and Dyer and Chernoff (1996). Herre, A. W. 1944. Notes on the fishes in the Zooo - Some of the character states in Notocheirus (includ - logical Museum of Stanford University. XVII. New ing lack of first dorsal, mesethmoid, first epibranchial fishes from Johore and India. Proceeding of the Bio - and epural) could be interpreted as paedomorphism logical Society of Washington, 57: 45-52. (eg. Weitzmann and Vari, 1988), and hence advanced Jordan, D. S. & C. L. Hubbs. 1919. Studies in Ichthy - rather than plesiomorphic traits. In any event, the dis - ology. A monographic review of the family tinction between Notocheiridae and Isonidae is sup - Atherinidae or silversides. Leland Stanford Junior ported by 20 characters. University Publications, University Series. 87 pp. Notocheirus is known only from coastal waters of Jordan, D. S. & E. C. Starks. (1901). A review of the South America, a region in which Iso has never been atherine fishes of Japan. Proceedings of the United collected. It is not known whether the separation States National Museum. 24: 198-206. aqua vol. 11 no. 1 - 2006 42 Basim Saeed, Walter Ivantsoff and Aarn

McCulloch, A. R. 1929. A check-list of the fishes recorded from Australia. Memoirs of the Australian Museum, 5 (1): 1-144. Ogilby, J. D . 1895. On two genera and species of fishes from Australia. Proceedings of the Linnean Society of N.S.W. 10: 320-324. Regan, C. T. 1919. fishes from Durban collected by Messrs. H. W. Bell Marley and Romer Robinson. Annals of Durban Museum, 2 (4): 197-204 . Rosen, D. E . (1964). The relationships and taxonomic position of the halfbeak, killifishes, silversides, and their relatives. Bulletin of the American Museum of Natural History, 127: 217-267. Saeed, B., Ivantsoff, W. & G. R. Allen . 1989. Taxo - nomic revision of the family Pseudomugilidae (Order Atheriniformes). Australian Journal of Marine and Freshwater Research, 40: 719-787. Saeed, B., Ivantsoff, W. & L. E. L. M. Crowley . 1993. A new species of the surf-inhabiting Atherini - form Iso (Pisces: Isonidae). Records of the Western Australian Museum. 16: 337-346. Saeed, B., Ivantsoff, W. & L. E. L. M. Crowley . 1994. Systematic relationships of atheriniform fami - lies within division 1 of the series Atherinomorpha (Acanthopterygii) with relevant historical perspec - tives. Journal of Ichthyology, 34 (9 ): 27-72. Said, B. M . 1983. Revision of the fish genus Iso . Unpublished MSc thesis, Macquarie University, Australia, 177 pp. Said, B . 1987. Revision of the genus Pseudomugil with phylogenetic systematics of the order Atherini - formes. Unpublished. Ph.D. thesis, Macquarie Uni - versity, Australia, 280 pp. Schultz, L. P . 1950. Correction for “A revision of six subfamilies of atherine fishes, with description of new genera and species”. Copeia, 1950: 150. Waite, E. R. 1904a. New records of recurrences of rare fishes from eastern Australia. No. 3. Records of the Australian Museum, 5: 231-244. Waite, E. R. 1904b. Synopsis of the fishes of New South Wales. Memoirs of the New South Wales Nat - uralist Club, 2:1-59. Weitzman, S. H. & R. P. Vari . 1988. Miniaturization in South American freshwater fishes; an overview and discussion. Proceedings of the Biological Society of Washington, 101: 444-465. Winterbottom, R . 1974. A descriptive synonymy of the striated muscles of the Teleostei. Proceedings of the Academy of Natural Sciences of Philadelphia, 125: 225-317.

43 aqua vol. 11 no. 1 - 2006 Index of aqua Vol. 10 (1-4) (Index by: 1. Author(s); 2. New Taxa; 3. Biology/Ecology/Biography/Reviews)

Author(s): Allen, Gerald R. and Mark V. Erdmann: Chromis xouthos , a new species of damselfish (Pomacentridae) from the East Andaman Sea and Central Indian Ocean. aqua 10 (3): 89-94, October 2005. Allen, Gerald R. and John E. Randall: A new species of damselfish (Pomacentrus : Pomacentridae) from Fiji. aqua 10 (3): 95-102, October 2005. Barreiros, João Pedro and Manuel Teves: The sunfish Mola mola as an attachment surface for the Lepadid Cirriped Lepas anatifera – a previously unreported association. aqua 10 (1): 1-4, June 2005. Herler, Jürgen and Helge Hilgers: A synopsis of coral and coral-rock associated gobies (Pisces: Gobiidae) from the Gulf of Aqaba, northern Red Sea. aqua 10 (3): 103-132, October 2005. Larson, Helen K., Ivantsoff, Walter and L. E. L. M. Crowley: Description of a new species of freshwater hardyhead, Craterocephalus laisapi (Pisces, Atherinidae) from East Timor. aqua 10 (2): 81-88, July 2005. Møller, Peter Rask, Werner Schwarzhans and Jørgen G. Nielsen : Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia . aqua 10 (4):133-205, November 2005. Motomura, Hiroyuki, Harazaki, Shigeru and Graham S. Hardy: A new species of triplefin (Perciformes: Tripterygiidae), Enneapterygius senoui, from Japan with a discussion of its in situ colour pattern. aqua 10 (1): 5-14, June 2005. Randall, John E.: Chlorurus perspicillatus x C. sordidus , a hybrid parrotfish from the Hawaiian Islands. aqua 10 (1): 39-43, June 2005. Randall, John E. and Gerald R. Allen: Neopomacentrus sororius , a new species of damselfish from the Indian Ocean, with description of a neotype for its sister species, N. azysron (Bleeker). aqua 10 (2): 73-80, July 2005. Schneidewind, Frank: An frogfish (Antennarius sp.) as a mimic of sea urchins: a new form of mimicry in the family Antennariidae. aqua 10 (1): 23-28, June 2005. Wetzel, James E., Poly, William J. and James W. Fetzner, Jr.: Orconectes pardalotus , a new species of crayfish (Decapoda: Cambaridae) from the lower Ohio River with notes on its life history. aqua 10 (2): 57-72, July 2005. Winterbottom, Richard: Two new species of the Trimma tevegae species group from the Western Pacific (Percomorpha: Gobiidae). aqua 10 (1): 29-38, June 2005. Winterbottom, Richard: Feia dabra, a new species of gobiid fish (Percomorpha: Gobiidae) from Palau. aqua 10 (2): 45-50, July 2005. Winterbottom, Richard: On the Status of Trimma tevegae and Trimma caudomaculata (Percomorpha: Gobiidae). aqua 10 (2):51-56, July 2005. Zuanon, Jansen and Ivan Sazima: The ogre catfish: prey scooping by the auchenipterid Asterophysus batrachus . aqua 10 (1): 15-22, June 2005.

New Taxa: Chromis xouthos n. sp. A new species of damselfish (Pomacentridae) from the East Andaman Sea and Central Indian Ocean. aqua 10 (3): 89-94, October 2005. Craterocephalus laisapi n. sp. Description of a new species of freshwater hardyhead, Craterocephalus laisapi (Pisces, Atherinidae) from East Timor. aqua 10 (2): 81-88, July 2005. Enneapterygius senoui n. sp. A new species of triplefin (Perciformes: Tripterygiidae), Enneapterygius senoui, from Japan with a discussion of its in situ colour pattern. aqua 10 (1): 5-14, June 2005. Feia dabra n. sp. a new species of gobiid fish (Percomorpha: Gobiidae) from Palau. aqua 10 (2): 45-50, July 2005. Neopomacentrus sororius n. sp. A new species of damselfish from the Indian Ocean, with description of a neotype for its sister species, N. azysron (Bleeker). aqua 10 (2): 73-80, July 2005. Ogilbia boehlkei n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia . aqua 10 (4):133-205, November 2005. Ogilbia boydwalkeri n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia . aqua 10 (4):133-205, November 2005. Ogilbia cocoensis n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia . aqua 10 (4):133-205, November 2005. Ogilbia davidsmithi n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia . aqua 10 (4):133-205, November 2005. Ogilbia jeffwilliamsi n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia . aqua 10 (4):133-205, November 2005. Ogilbia jewettae n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia . aqua 10 (4):133-205, November 2005. Ogilbia mccoskeri n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia . aqua 10 (4):133-205, November 2005. Ogilbia nigromarginata n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia . aqua 10 (4):133-205, November 2005. Ogilbia nudiceps n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia . aqua 10 (4):133-205, November 2005. Ogilbia robertsoni n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia . aqua 10 (4):133-205, November 2005. Ogilbia sabaji n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia . aqua 10 (4):133-205, November 2005. Ogilbia sedorae n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia . aqua 10 (4):133-205, November 2005. Ogilbia suarezae n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia . aqua 10 (4):133-205, November 2005. Orconectes pardalotus n. sp. A new species of crayfish (Decapoda: Cambaridae) from the lower Ohio River with notes on its life history. aqua 10 (2): 57-72, July 2005. Ogilbia tyleri n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia . aqua 10 (4):133-205, November 2005. Pomacentrus microspilus n. sp. A new species of damselfish (Pomacentrus : Pomacentridae) from Fiji. aqua 10 (3): 95-102, October 2005. Trimma marinae n sp. Two new species of the Trimma tevegae species group from the Western Pacific (Percomorpha: Gobiidae). aqua 10 (1): 29-38, June 2005. Trimma nasa n sp. Two new species of the Trimma tevegae species group from the Western Pacific (Percomorpha: Gobiidae). aqua 10 (1): 29-38, June 2005.

Biology/Ecology/Biography/Reviews: A frogfish (Antennarius sp.) as a mimic of sea urchins: a new form of mimicry in the family Antennariidae. aqua 10 (1): 23-28, June 2005. A synopsis of coral and coral-rock associated gobies (Pisces: Gobiidae) from the Gulf of Aqaba, northern Red Sea. aqua 10 (3): 103-132, Octo - ber 2005. Book review: Freshwater Fishes of north-eastern Australia. aqua 10 (4): 208, November 2005. Book review: Reef and Shore Fishes of the South Pacific New Caledonia to Tahiti and the Pitcairn Islands. aqua 10 (3):102, October 2005. Chlorurus perspicillatus x C. sordidus , a hybrid parrotfish from the Hawaiian Islands. aqua 10 (1): 39-43, June 2005. On the Status of Trimma tevegae and Trimma caudomaculata (Percomorpha: Gobiidae). aqua 10 (2):51-56, July 2005. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia . aqua 10 (4):133-205, November 2005. The ogre catfish: prey scooping by the auchenipterid Asterophysus batrachus . aqua 10 (1): 15-22, June 2005. The sunfish Mola mola as an attachment surface for the Lepadid Cirriped Lepas anatifera – a previously unreported association. aqua 10 (1): 1-4, June 2005.

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References to literature : In both the text and the reference such, ideally in parentheses and including the initials of the (senior) list, the name-year system must be used. The list of references author, eg "(Note: Fig. 1 here. AZ)". should be placed at the end of the paper, alphabetically arranged 10. Evaluation of manuscripts: manuscripts submitted to aqua according to author name. Only those publications cited in the will be evaluated by the editors and referees on the basis of their paper may be included. Titles of journals should be given in full. contents. Papers are accepted on the understanding that they Examples of correct reference formats: have not and will not be published elsewhere. Blaber, S. J. M. 1980. Fish of the Trinity Inlet System of North 11. Reprints: Authors will receive 1 free reprint and 1 PDF-file of Queensland, with Notes on the Ecology of Fish Fauna of Tropic- each paper . Additional reprints may be ordered from Aquapress. aqua Journal of Ichthyology and Aquatic Biology Vol. 11 (1), February 2006 Contents: Gerald R. Allen: Cirrhilabrus brunneus , a new wrasse (Pisces: Labridae) from north-eastern Kalimantan, Indonesia ...... 1-4 Wilson J. E. M. Costa: Redescription of Kryptolebias ocellatus (Hensel) and K. caudomarginatus (Seegers) (Teleostei: Cyprinodontiformes: Rivulidae), two killifishes from mangroves of south-eastern Brazil ...... 5-12

Gerald R. Allen, Forrest Young and Patrick L. Colin: Centropyge abei , a new species of deep-dwelling angelfish (Pomacanthidae) from Sulawesi, Indonesia ...... 13-18 John E. Randall and John L. Earle: Amblyeleotris neumanni, a new species of shrimp goby from New Britain ...... 19-24 Basim Saeed, Walter Ivantsoff and Aarn: Descriptive anatomy of Iso rhothophilus (Ogilby), with a phylogenetic analysis of Iso and a redefinition of Isonidae (Atheriniformes) ...... 25-43

Index aqua 10(1-4) ...... 44

Papers appearing in this journal are indexed in: Zoological Record; Biolis - Biologische Literatur Information Senckenberg; www.aquapress-bleher.it; www.aquageo.com; www.Joachim-Frische.com

Cover photo: Kryptolebias ocellatus , hermaphrodite, UFRJ 6243, 46.8 mm SL; Brazil: Rio de Janeiro: Guaratiba. Photo by W. J. E. M. Costa.

Male and female of the wild common carp from the spawning school at the Lesser Danube above Kolárovo (1955). Photo from the forthcoming Ms “ The oldest domesticated fishes, and the consequences of an epigenetic dichotomy in fish cul - ture” in aqua Vol. 11 (2). Photo by E. K. Balon.