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

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

3 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: Keywords Taxonomy, marine fishes, Cirrhilabrus, new species, Labridae, Kalimantan, Indonesia Abstract Cirrhilabrus brunneus is described from a single male specimen, 43.6 mm SL, collected at north-eastern Kalimantan, Indonesia. It is one of only three species in the genus that possesses a lunate caudal fin. The new species, which is overall dark brown in colour, including the median and pelvic fins, closely resembles C. lunatus from the Ryukyu Islands, southern Japan, and the Ogasawara Islands. However, unlike the latter species, it lacks a broad orange-yellow zone along the sides at the level of the pectoral fins, a pale yellowish band along the base of the dorsal fin, and irregular diagonal red lines on the snout and the postorbital portion of the head that extends onto the anterodorsal part of the body. There is also a substantial difference in maximum size with males of C. lunatus attaining a maximum standard length of about 85 mm in comparison with less than 50 mm for C. brunneus. Introduction The labrid genus Cirrhilabrus Temminck and Schlegel contains small, colourful, sexually dimorphic fishes that inhabit Indo-west Pacific coral reefs. Allen (2000) listed 36 species, but since then six additional species have been described by Senou and Hirata (2000), Randall and Pyle (2001), Randall and Nagareda (2002), and Allen et al. (2003). The members of the genus typically live in rubble habitats, frequently below 20 m, and feed on zooplankton well above the bottom. Due to their relatively deepdwelling habits, most species remained undetected until the widespread use of scuba equipment by scientific divers over the past few decades. No doubt additional discoveries will occur in the future, especially with the increased use of specialized deep diving equipment such as mixed-gas scuba and small research submarines. The present paper describes a new species that was collected by the author during a recent coral reef survey for The Nature Conservancy at the Berau district of north-eastern Kalimantan, Indonesia. Materials and Methods Counts of fin spines are given in Roman numerals and soft rays in Arabic. Pectoral ray counts include the rudimentary upper ray. The lateral line is interrupted; the count of the anterior part is given first, followed by a plus sign and the peduncular part. Only lateral line scales with tubes are counted. All the tubed scales of the peduncular part are counted, even though one is usually located posteriorly to the base of the caudal fin. The number of scales in the rows on the cheek is counted from where they commence below the front of the orbit to behind the centre of the orbit. Gill raker counts include all rudiments. Because it may be difficult to determine which raker is at the angle, only the total gill raker count is given. Lengths of specimens are given as standard length (SL) except estimates of total length (TL) of fishes photographed underwater; this is the straight-line measurement from the front of the upper lip to the base of the caudal fin (end of hypural plate). Measurements are given as percentages of the standard length. Head length is the distance from the front of the upper lip to the posterior end of the opercular membrane. Body depth is the greatest depth to the base of the dorsal fin (adjusting for any malformation of the abdomen due to preservation). Body width is measured just posterior to the opercular flap. Snout length is taken from the front of the upper lip to the fleshy edge of the orbit (if the upper jaw is protruded, it is pressed back to the non-protractile position before the measurement is taken. The same is true of SL and head length measurements. Orbit diameter is the greatest fleshy diameter. Interorbital width is the least bony width. Caudal peduncle depth is the least depth; caudal peduncle length is the horizontal measurement between verticals at the rear base of the anal fin and the caudal fin base. Measurements of fin spines and rays are taken to the extreme base of these elements. Caudal concavity is the horizontal distance between the tips of the longest and shortest caudal rays. Pectoral fin length is taken from the tip of the longest ray to the base of this ray. Pelvic fin length is measured from the base of the spine to the tip of the longest ray. The holotype and only known specimen is deposited at Pusat Penelitian dan Pengembangan Oseanologi, Jakarta, Indonesia (NCIP). 1 aqua vol. 11 no

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

5 Gerald R. Allen elongate scales, one per membrane (except first scale which covers membranes of first and second spines), the longest about one-half spine length (basal scales progressively shorter posteriorly on membranes of soft portion of fin); peduncular lateral line scales followed by one slightly larger pored scale (included in lateral line count) on base of caudal fin with a slightly posterior scale above and below, these three scales followed by a vertical row of three large scales, the middle one overlapping the ones above and below, reaching two-thirds distance to posterior margin of fin; pectoral fins scaleless; pelvic fins with a median ventral process of two elongate scales, the more pointed posterior scale extending to tip of depressed pelvic spine spine; slender axillary scale of each pelvic fin extending about equal in length to pelvic spine. Origin of dorsal fin above third lateral line scale; first dorsal spine 4.8 (7.1) in head; remaining dorsal spines progressively longer, the last 1.9 (18.1) in head; interspinous membranes of dorsal fin extending above spine tips, supported by the terminal cirrus projecting upward from just behind each spine tip; first to third dorsal soft rays the longest, 2.1 in head; origin of anal fin on a vertical with base of penultimate dorsal spine; first anal spine 4.5 (7.6) in head; second anal spine 3.9 (8.7) in head; third anal spine 3.4 (10.1) in head; fourth and fifth anal soft rays longest, 2.4 (14.2) in head; caudal fin lunate with protruding upper and lower lobes, its length 1.1 (30.3) in head; caudal concavity 2.4 (14.2) in head; third and fourth pectoral rays longest, 1.5 (22.9) in head; origin of pelvic fins below pectoral in base; pelvic fin length 1.5 (22.7) in head, the pelvic fin tips falling well short of anal fin origin when depressed; length of pelvic fin spine 3.0 (11.2) in head. Colour of male holotype when fresh (Fig. 1): mainly dark brown with slight purplish hue; breast and belly with bronze hue, a diamond shaped, red-brown marking on each scale; iris bright red and pupil blackish; dorsal fin dark brown with white distal margin that gradually widens posteriorly, resulting in a broad white area around angular posteriormost portion of fin; anal fin dark brown with narrow white margin; caudal fin mainly brown including prominent upper and lower margins of fin, the posterior margin broadly white; pelvic fins dark brown; pectoral rays translucent with dark brown base. Colour or male holotype in alcohol: after more than two years of preservation, the colour pattern is remarkably similar to that described in the previous paragraph with the exception of a tan iris and tannish lower onethird of the body. Live colour of female from underwater field notes: overall brownish (red when illuminated with underwater torchlight) with faint longitudinal dark stripes on head and body and small dark spot on upper caudal peduncle. Remarks The caudal fin shape of the male serves to separate C. brunneus from nearly all other members of the genus which generally have truncate, emarginate, rhomboid, or lanceolate caudal fins. The only other species with produced fin lobes are C. lunatus Randall and Masuda (1991) from the Ryukyu Islands, southern Japan, and the Ogasawara Islands, and C. johnsoni Randall (1988) from the Marshall Islands. Males of both species have strongly lunate caudal fins, with the upper and lower rays forming filamentous extensions. The distinctive colour patterns of the three species with lunate-shaped caudal fins are diagnostic. The male of C. johnsoni is primarily orange-red with bright red median fins. The new species appears to be most closely related to C. lunatus (Figure 2). The two species have a similar dorsal fin shape and there are remarkable similarities in the coloration of the median fins, which are dark (brown in C. brunneus and black in C. lunatus) with a broad white margin posteriorly on the dorsal and caudal fins. However, C. lunatus differs in having a broad orange-yellow zone along the sides at the level of the pectoral fins and fish from the Ryukyu Islands have a purplish coloration on the sides, which angles upward posteriorly to encompass the entire caudal peduncle. Male specimens of C. lunulatus from the Ogasawara Islands have pinkish coloration on the lower sides. Males from both the Ryukyu and Ogasawara islands differ from those of C. brunneus in having a pale yellowish band along the base of the dorsal fin and irregular diagonal red lines on the snout and postorbital portion of the head that extend onto the anterodorsal part of the body. There is also a possible difference in the number of gill Fig. 2. Cirrhilabrus lunatus, freshly collected male holotype, 67.7 mm SL, Okinawa, Ryukyu Islands. Photo by J. E. Randall. Fig. 3. Cirrhilabrus lunatus, freshly collected female paratype, 45.1 mm SL, Ani-jima, Ogaswara Islands. Photo by J. E. Randall. 3 aqua vol. 11 no

6 Cirrhilabrus brunneus, a new wrasse (Pisces: Labridae) from north-eastern Kalimantan, Indonesia rakers on the first branchial arch. Randall and Masuda (1991) indicated a range of gill rakers for six specimens of C. lunatus in comparison to 13 rakers for the holotype of C. brunneus. Of course, it would be desirable to count additional specimens of the latter species to verify this disparity. There is also a substantial difference in maximum size. Males of C. lunatus attain a maximum standard length of at least 84.9 mm. In comparison, the holotype of C. brunneus, measuring 43.6 mm SL, is scarcely more than half this size. Moreover, four more males of similar size were sighted at the type locality. Several initial phase (female) individuals of C. brunneus were observed at the type locality. They were very similar in appearance to the female of C. lunatus from the Ogasawara Islands that was illustrated by Randall and Masuda (1991) and is included here as Fig. 3. Cirrhilabrus brunneus is presently known only from the 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. from Fiji. aqua, Journal of Ichthyology and Aquatic Biology, 7 (3): Randall, J. E Five new wrasses of the genera Cirrhilabrus and Paracheilinus (Perciformes: Labridae) from the Marshall Islands. Micronesica, 21: Randall, J. E. & H. Masuda, Two new labrid fishes of the genus Cirrhilabrus from Japan. Revue française d Aquariologie, 18 (2): Randall, J.E. & B.H. Nagareda Cirrhilabrus bathyphilus, a new deep-dwelling labrid fish from the Coral Sea. Cybium, 26 (2): Randall, J.E. & R.L. Pyle Three new species of labrid fishes of the genus Cirrhilabrus from islands of the tropical Pacific. aqua, Journal of Ichthyology and Aquatic Biology, 4 (3): Senou, H. and T. Hirata A new labrid fish, Cirrhilabrus katoi, from southern Japan. Ichthyological Research, 4 (1): Etymology This species is named brunneus (Latin: dusky or dark) with reference to the overall colour pattern of the holotype. 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 Ecological 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 Description of a new wrasse (Pisces: Labridae: Cirrhilabrus) from northern Sumatra, Indonesia. aqua, Journal of Ichthyology and Aquatic Biology, 4 (2): Allen, G. R., J. E. Randall & B. A. Carlson Cirrhilabrus marjorie, a new wrasse (Pisces: Labridae) aqua vol. 11 no

7 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 , Rio de Janeiro, Brasil. wcosta@acd.ufrj.br Accepted: Keywords Fish, Cyprinodontiformes, Rivulidae, Kryptolebias, Neotropica, mangrove, systematics, hermaphrodite, morphology Abstract Taxonomy of Kryptolebias ocellatus (Hensel) and K. caudomarginatus (Seegers) has been poorly defined and mostly based on a few specimens bred in aquaria. They are considered valid species and are redescribed based on recent collections in the type locality area, the mangroves of Rio de Janeiro state, south-eastern Brazil. Both species are considered closely related to K. marmoratus (Poey), with which they share the presence of four neuromasts on the posterior supraorbital series, a long anterior nostril, and a bony laminar ventral process on the fifth ceratobranchial; K. ocellatus is distinguished by a unique colour pattern in the hermaphrodite, more slender caudal peduncle, and a shorter pelvic fin, and K. caudomarginatus by possessing more vomerine teeth, longer dorsal fin base, and a unique male colour pattern. The type locality of K. caudomarginatus is coorrected. Resumo A taxonomia de Kryptolebias ocellatus (Hensel) e K. caudomarginatus (Seegers) tem sido fracamente definida, principalmente com base em poucos espécimes criados em aquários. Elas são consideradas espécies válidas e são redescritas com base em coleções recentes na área da localidade tipo, os manguezais do estado do Rio de Janeiro, sudeste do Brasil. Ambas espécies são consideradas estreitamente aparentadas a K. marmoratus (Poey), com a qual elas compartilham a presença de quatro neuromastos na série supraorbital posterior, narina anterior longa, e processo ventral laminar ósseo em quinto ceratobranquial; K. ocellatus é distinguível por possuir um padrão de colorido exclusivo em hermafrodita, pedúnculo caudal mais esguio e nadadeira pélvica mais curta, e K. caudomarginatus por possuir mais dentes no vomer, base de nadadeira dorsal mais longa e padrões de colorido exclusivos em macho. A localidade tipo de K. caudomarginatus é corrigida. Zusammenfassung Die systematische Stellung von Kryptolebias ocellatus (Hensel) und K. caudomarginatus (Seegers) wurde bisher nicht ausreichend begründet und ihre Beschreibung beruhte auf einigen wenigen Exemplaren aus Aquarien. Sie werden hier zu gültigen Arten erklärt und auf der Grundlage neuerer Fänge im Gebiet der Typuslokalität neu beschrieben, den Mangrovensümpfen der Region Rio de Janeiro, in SO-Brasilien. Die beiden Arten werden als nahe verwandt mit K. marmoratus (Poey) eingestuft; gemeinsame Merkmale sind vier Neuromasten in der hinteren supraorbitalen Reihe, ein langes vorderes Nasenloch und ein knochiger, blättchenhafter ventraler Fortsatz am fünften Ceratobranchiale; K. ocellatus unterscheidet sich durch ein unverkennbares Farbmuster ohne Geschlechtsunterschied, einen schlankeren Schwanzstiel und eine kürzere Bauchflosse, während K. caudomarginatus mehr Schlundzähne, eine längere Rückenflossenbasis und ein unverkennbares männliches Farbmuster besitzt. Die Typuslokalität von K. caudomarginatus wird korrekt abgegrenzt. Résumé La taxinomie des espèces Kryptolebias ocellatus (Hensel) et K. caudomarginatus (Seegers) a été peu spécifiée et basée surtout sur un petit nombre de spécimens élevés en aquarium. Elles sont considérées comme des espèces valides et sont redécrites sur base de collectes récentes dans la région de la localité-type, les mangroves de l'état de Rio de Janeiro, au sud-est du Brésil. Les deux espèces paraissent très proches de K. marmoratus (Poey), et, comme celle-ci, montrent la présence de quatre neuromastes sur la série postérieure supra-orbitale, une longue narine antérieure et une excroissance osseuse laminaire ventrale sur la cinquième cératobranchiale. K. ocellatus se distingue par une coloration unique du pédoncule caudal hermaphrodite, plus mince et une pelvienne plus courte, et 5 aqua vol. 11 no

8 Redescription of Kryptolebias ocellatus (Hensel) and K. caudomarginatus (Seegers) (Teleostei: Cyprinodontiformes: Rivulidae) K. caudomarginatus, par un plus grand nombre de dents vomériennes, une plus longue base de la dorsale et un patron de coloration unique pou le mâle. La localité-type de K. caudomarginatus est correctement identifiée. Sommario La tassonomia di Kryptolebias ocellatus (Hensel) e K. caudomarginatus (Seegers) è poco definita e basata per lo più su pochi esemplari provenienti da accoppiamenti in acquario. Esse sono considerate specie valide e vengono qui descritte nuovamente sulla base di recenti catture nella località tipo, le mangrovie dello stato di Rio de Janeiro, Brasile sud-orientale. Entrambe sono considerate molto vicine a K. marmoratus (Poey), con cui condividono la presenza di quattro neuromasti sulla serie supraorbitale posteriore, una lunga narice anteriore e un processo laminare ventrale osseo sul quinto ceratobrachiale; K. ocellatus si distingue per una tipica colorazione nell ermafrodita, un peduncolo caudale più sottile e le pinne pelviche più corte, mentre K. caudomarginatus si differenzia per possedere più denti vomerini, una pinna dorsale a base più lunga e una singolare colorazione nel maschio. La località tipo K. caudomarginatus è definita in modo corretto. Introduction Kryptolebias Costa was recently established to include some species previously placed in Rivulus: K. brasiliensis (Valenciennes), K. caudomarginatus (Seegers), and K. ocellatus (Hensel) from south-eastern Brazil; K. campelloi (Costa) from eastern Brazilian Amazon; K. marmoratus (Poey), recorded from a broad geographic range, including the south-eastern United States, Bahamas, Yucatan Peninsula, Cuba and other Caribbean islands, Venezuela and Guianas (Costa, 2004a, 2004b). Three poorly known nominal species, considered synonyms of K. marmoratus (e. g., Costa, 2003), were also included in Kryptolebias: K. bonairensis (Hoedeman) from Bonaire, Curação, and coastal Venezuela; K. heyei (Nichols) from Isla Saona, Dominican Republic; and, K. garciai (De la Cruz & Dubitsky) from Cuba (Costa, 2004a). Subsequently, Vermeulen & Hrbek (2005) described K. sepia from Surinam. Kryptolebias has been hypothesized to be the sister group to a clade comprising all other rivulids on the basis of morphological and some molecular analyses (Hrbek & Larson, 1999; Costa, 2004a, 2004c; Vermeulen & Hrbek, 2005) or a basal clade within a monophyletic lineage including all rivulids except cynolebiatines according to other molecular analyses (Murphy et al., 1999; Vermeulen & Hrbek, 2005). Kryptolebias brasiliensis, K. campelloi, and K. sepia are similar to most rivulids in inhabiting freshwater pools and streams (e. g., Costa, 2004a; Vermeulen & Hrbek, 2005), while K. caudomarginatus, K. ocellatus, and K. marmoratus live in brackish to salt water, usually in mangrove swamps (e. g., Taylor, 1988; the present paper). Both morphological (Costa, 2004a) and molecular studies (Murphy et al., 1999; Vermeulen & Hrbek, 2005) support monophyly of the latter assemblage. Among species of this assemblage, K. ocellatus and K. marmoratus are self-fertilizing hermaphroditic species, a condition unique among vertebrates (e. g., Harrington, 1961). Hermaphrodites are female-like individuals, exhibiting typical female colour patterns. Populations are mainly composed of hermaphrodites, while functional females are unknown, and males are uncommon or very rare (e. g., Davis et al., 1990; Turner et al., 1992). For example, Taylor (1990) collected over 250 specimens of K. marmoratus in Florida, but none of them was a male. Although some species of Kryptolebias are among the oldest described species of the Rivulidae, their taxonomy is still confused. The best example is K. ocellatus, described by Hensel (1868) based on a single specimen collected in Rio de Janeiro. The identity of K. ocellatus was difficult to determine between 1906 and 1984, a period when K. caudomarginatus, another species from Rio de Janeiro and undescribed at that time, was usually identified as Rivulus ocellatus (= K. ocellatus). Seegers (1984) established the identity of K. ocellatus and first described K. caudomarginatus based upon aquarium bred specimens. Seegers also considered K. marmoratus conspecific to K. ocellatus, proposing the two names as subspecies of K. ocellatus, which would have chronological priority over K. marmoratus, described 12 years later (Poey, 1980). However, K. marmoratus is an important experimental fish for researches on carcinogenicity, mutagenesis, teratogenesis, and other scientific areas, in which the specific epithet marmoratus is consistently applied, whereas the specific epithet ocellatus, the putative senior synonym of marmoratus, was poorly known and never correctly applied for many decades. These were the basic arguments used by Lazara & Smith (1990) to propose suppression of the specific epithet ocellatus and conservation of the name marmoratus. However, Costa (1994) did not agree with such a proposal, which would be premature, since no rigorous taxonomic study was developed to check whether ocellatus and marmoratus were in fact the same species. The International Commission of Zoological Nomenclature (1995) then decided to give precedence to the name marmoratus over the name ocellatus, whenever the two names are considered to be synonyms. However, due to difficulties in collecting small fishes in a mangrove habitat, K. ocellatus and K. caudomarginatus have been insufficiently sampled in scientific collections, and consequently are still poorly known taxonomically. Both species are herein redescribed, based on recent collections made in their type locality areas. Material and methods All material was collected with dip nets. Measurements and counts follow Costa (1995). Measurements are pre- aqua vol. 11 no

9 Wilson J. E. M. Costa sented as percentages of standard length (SL), except those related to head morphology, and are expressed as percentages of head length. Fin ray counts include all elements. Numbers of vertebrae, gill rakers, and pectoral, pelvic and caudal fin rays were recorded only from cleared and stained specimens. The compound caudal centrum was counted as a single element. Osteological preparations were made according to Taylor and Van Dyke (1985). Terminology for frontal squamation and the cephalic neuromast series follow Hoedeman (1958) and Costa (2001), respectively. The abbreviation c&s stands for specimens cleared and stained for bone and cartilage. Material is deposited in Universidade Federal do Rio de Janeiro, Rio de Janeiro, Brazil (UFRJ). Other abbreviations for of institution names are: ZFMK, Zoologisches Farschungsinstitut und Museum Alexander Koenig, Bonn, and ZMB, Zoologisches Museum, Humboldt-Universität, Berlin. Kryptolebias ocellatus (Hensel) (Figs. 1-2; Table I) Rivulus ocellatus Hensel, 1868: 365 (type locality: Rio de Janeiro; holotype: ZMB 7448). Material examined Brazil: Estado do Rio de Janeiro: UFRJ 1806, 3; mangrove swamp near Itaguaí; M. S. Melgaço, 25 Jul UFRJ 6234, 4 (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, UFRJ 6237, 3; idem; W. J. E. M. Costa, C. P. Bove & T. Litz, 19 Feb UFRJ 6243, 3; idem; W. J. E. M. Costa, J. Barata, A. Moreira & J. P. Moreira, 14 Aug UFRJ 6252, 9; idem; W. J. E. M. Costa, J.L. Mattos, L. Villa Verde, F. Ottoni & E. Mattos, 22 Oct Material collected in mangrove swamp near Caxias, baía de Guanabara, was not preserved. Diagnosis Distinguished from all rivulids except K. marmoratus and K. caudomarginatus in having long anterior nostril, its tip extending beyond anterior profile of the head. Also similar to K. marmoratus and K. caudomarginatus by possessing numerous scales in the longitudinal series (44-48, vs K. brasiliensis and K. campelloi, and in K. sepia) and four neuromasts on the posterior supraorbital series (vs. three). It differs from K. marmoratus and K. caudomarginatus in having a vertical row of pale silver to pale gold dots on the dorsal portion of flank in the hermaphrodite (vs. pale silver spots over the whole flank in the hermaphrodite of K. marmoratus and in the female of K. caudomarginatus), more slender caudal peduncle (caudal peduncle depth % SL, vs % SL), and shorter pelvic fin (pelvic fin length % SL, vs % SL). Description The description is based on hermaphrodites only. Morphometric data appear in Table I. Largest specimen 49.8 mm SL. Dorsal profile approximately straight to slightly convex from snout to end of dorsal fin base, approximately straight on caudal peduncle. Ventral profile gently convex from lower jaw to end of anal fin base, about straight on caudal peduncle. Body slender, subcylindrical anteriorly, about as deep as wide, to compressed posteriorly. Greatest body depth at vertical just in front to pelvic fin base. Jaws short, snout blunt. Anterior naris cylindrical and long, extending beyond anterior profile of the head. Urogenital papilla pocket-shaped. 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 reaching vertical just posterior to midlength between pectoral fin and pelvic fin bases. Pelvic fin elliptical, tip reaching base of 1st anal fin ray. Pelvic fin bases medially separated by short interspace. Dorsal fin origin on vertical between base of 10th and 11th anal fin rays, and 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

10 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) Percent of standard length Body depth Caudal peduncle depth Predorsal length Prepelvic length Length of dorsal fin base Length of anal fin base Caudal fin length Pectoral fin length Pelvic fin length Head length Percent of head length Head depth Head width Snout length Lower jaw length Eye diameter Fig. 2. Kryptolebias ocellatus, hermaphrodite, UFRJ 6237, 36.8 mm SL; Brazil: Rio de Janeiro: Guaratiba. Photo by W. J. E. M. Costa. between neural spines of 18th and 20th vertebrae. Anal fin origin between pleural ribs of 14th and 15th vertebrae. 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 squamation 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 without exposed margins. Longitudinal series of scales 47-48; transverse series of scales 13-14; scale rows around caudal peduncle Contact organs absent. Cephalic neuromasts: supraorbital 3 + 4, parietal 1-2, anterior rostral 1, posterior rostral 1, infraorbital , preorbital 3, otic 1, post-otic 2, supratemporal 1, median opercular 1, ventral opercular 2, preopercular , mandibular , 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 Vomerine teeth 1-4. Ventral process of posttemporal long. Total vertebrae 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. Dorsum light brown. Venter white. Side of head light aqua vol. 11 no

11 Wilson J. E. M. Costa brown on dorsal portion, pale silver with great concentration on melanophores ventrally. Lower jaw grey. Iris brown. Unpaired fins grey with hyaline small spots on basal portion; anal fin with pale yellow subdistal stripe, and distal grey stripe. Pelvic and pectoral fins hyaline. Distribution and habitat Kryptolebias ocellatus inhabits the shallow parts (about cm deep) of mangrove swamps associated with Sepetiba and Guanabara bays (Fig. 3). It is always sympatric to K. caudomarginatus, which is conspicuously more abundant, and sometimes with two species of poeciliids, Poecilia vivipara Bloch & Schneider and Phalloptychus januarius (Hensel), an eleotridine gobiid, Dormitator maculatus (Bloch), and the callichthyid Callichthys callichthys (Linnaeus). It is never found in freshwater streams. The occurrence of this species has already been reported in Santos, São Paulo (Huber, 1992), but the material from this region was not examined in the present paper. Remarks All the material herein examined included only hermaphrodites. Three recent collecting trips were unsuccessful in collecting males. However, Notare (1988) provided a colour photo of a live male identified as Rivulus Fig. 3. Mangrove at Guaratiba, Rio de Janeiro, Brazil, habitat of Kryptolebias ocellatus and K. caudomarginatus. Photo by W. J. E. M. Costa. caudomarginatus var. (?) collected in Rio de Janeiro and Huber (1992) illustrated a similar live male, also from Rio de Janeiro, identified as primary male of ocellatus. Notare s colour photo shows a fish in which the flank is light grey with dark red spots. There is a dark grey humeral spot and a grey caudal peduncle spot, and, the unpaired fins have a pale yellow subdistal zone and a dark grey to black distal zone. This material was not preserved for study, and its identification needs confirmation. Kryptolebias caudomarginatus (Seegers) (Figs. 4-7; Table I) Rivulus caudomarginatus Seegers, 1984: 307 (type locality: Greta Funda [correctly Grota Funda, an equivocal type locality as discussed in Remarks below] near a technical centre of the Army, southern Rio de Janeiro [correctly near Technological Center of the Brazilian Army, Guaratiba, south-western Município do Rio de Janeiro, Estado do Rio de Janeiro, southeastern Brazil, rio Piracão basin, a tributary to Baía de Sepetiba, S W, altitude 8 m]; holotype: ZFMK 12848). Material examined Brazil: Estado do Rio de Janeiro: UFRJ 1805, 32; mangrove near Itaguaí; M.S. Melgaço, 25 Jul UFRJ 1047, 22; UFRJ 3583, 3 (c&s); Madeira, mangrove swamp or mangrove habitat near Itaguaí; M. S. Melgaço, 25 Jul 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 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, UFRJ 6238, 4; idem; W. J. E. M. Costa, C. P. Bove & T. Litz, 19 Feb UFRJ 6244, 12; idem; W. J. E. M. Costa, J. Barata, A. Moreira & J. P. Moreira, 14 Aug UFRJ 6253, 4; idem; W. J. E. M. Costa, J. L. Mattos, L. Villa Verde, F. Ottoni & E. Mattos, 22 Oct Material collected in mangrove habitat 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 similar to K. marmoratus and K. ocellatus by possessing numerous scales in the longitudinal series (44-48, vs K. brasiliensis and K. campelloi, and in K. sepia in other species of Kryptolebias) and four neuromasts 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 ( % SL, vs % SL), and a unique colour pattern of flank and caudal fin in male (flank with black oblique bars on its posterior half, vs. dark red 9 aqua vol. 11 no

12 Redescription of Kryptolebias ocellatus (Hensel) and K. caudomarginatus (Seegers) (Teleostei: Cyprinodontiformes: Rivulidae) Fig. 4. Kryptolebias caudomarginatus, male, UFRJ 6235, 34.8 mm SL; Brazil: Rio de Janeiro: Guaratiba. Photo by W. J. E. M. Costa. Pelvic fin bases medially separated by short interspace or in contact. Dorsal fin origin on vertical between base 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; pectoral-fin rays 13; pelvic fin rays 5-6. Scales small, cycloid. Body and head entirely scaled, except anterior ventral surface of head. Body squamation 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 without exposed margins. Longitudinal series of scales 43-48; transverse series of scales 9-10; scale rows around caudal peduncle Contact organs absent. Cephalic neuromasts: supraorbital , parietal 2, anterior rostral 1, posterior rostral 1, infraorbital , preorbital 3, otic 1, postotic 2, supratemporal 1, median opercular 1, ventral opercular 2, preopercular , 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 % basihyal length. Six branchiostegal rays. Second pharyngobranchial teeth 3-5. Gill rakers of first branchial arch 1-2 Fig. 5. Kryptolebias caudomarginatus, male, UFRJ 6238, 42.4 mm SL; Brazil: Rio de Janeiro: Guaratiba. Photo by W. J. E. M. Costa. spots; caudal fin with a broad white to yellowish white subdistal zone, strongly contrasting with a broad black distal zone, vs. indistinct pale yellow subdistal zone bordered 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, approximately straight on caudal peduncle. Ventral profile gently convex from lower jaw to end of anal fin base, about straight on caudal peduncle. Body slender, subcylindrical anteriorly, about as deep as wide, to compressed posteriorly. Greatest body depth at vertical just in front of pelvic fin base. Jaws short, snout blunt. Anterior naris cylindrical and long, extending anteriorly beyond anterior 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 reaching 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. 6. Kryptolebias caudomarginatus, female, UFRJ 6235, 31.9 mm SL; Brazil: Rio de Janeiro: Guaratiba. Photo by W. J. E. M. Costa. Fig. 7. Kryptolebias caudomarginatus, female, UFRJ 6238, 36.3 mm SL; Brazil: Rio de Janeiro: Guaratiba. Photo by W. J. E. M. Costa. aqua vol. 11 no

13 Wilson J. E. M. Costa Vomerine teeth Ventral process of posttemporal long. Total vertebrae Coloration in life: Male: Side of body pale bluish silver; black spots irregularly shaped and arranged on anterior half of flank and black oblique bars, usually zigzag shaped, sometimes broken, forming isolated black spots on posterior half of flank; approximately rectangular black humeral blotch; round black spot on dorsal end of caudal peduncle, sometimes other black spot below. Dorsum light brown, with green iridescence of laterodorsal scales. Venter white. Side of head light greenish brown on dorsal portion, pale silver with great concentration of melanophores ventrally. Lower jaw grey. Iris brown. Dorsal and anal fins light grey with small dark grey to black small spots, broad subdistal white stripe and narrow distal black stripe. Caudal fin dark grey, with broad white to yellowish-white subdistal zone and broad black distal zone; often dorsal and ventral margins and filaments dark orange. Pelvic fin hyaline with subdistal white line and black margin. Pectoral fin hyaline. Female: Side of body light brown with pale silver spots; black humeral blotch; round black spot with bright grey margin on dorsal end of caudal peduncle, Caudal fin hyaline, with small faint grey spots on basal portion and pale grey margin. Pelvic and pectoral fins hyaline. All described color patterns visible in preserved specimens, both in male and female. Distribution and habitat As described for K. ocellatus. Remarks During the 1980 s, several collections of K. ocellatus and K. caudomarginatus were made in a mangrove area near the Technological Center of the Brazilian Army, Guaratiba, south-western part of the metropolitan region of the Rio de Janeiro city, by aquarists. Seegers (1984) mentioned that the type specimens of R. caudomarginatus were the progeny of fish collected in Greta Funda, a misspelling of Grota Funda, near a technical center of the Army. However, Grota Funda is situated about 4 km east of the Technological Center of the Brazilian Army, at a small coastal hill (Serra Geral de Guaratiba), at an altitude of m. No habitat suitable to the occurrence of rivulids is present in Grota Funda, thus it is considered an equivocal type locality. Discussion Monophyly of the assemblage including K. ocellatus, K. marmoratus and K. caudomarginatus has been supported by different studies (e. g., Murphy et al., 1999; Costa, 2004a; Vermeulen & Hrbek, 2005). New evidence corroborating this hypothesis is herein provided: 1) presence of four neuromasts on the posterior supraorbital series (Fig. 8), a condition not occurring in other species of Kryptolebias nor in members of other basal rivulid lineages, such as the genera Prorivulus Costa and Rivulus Poey, which have three neuromasts; 2) long anterior nostril (Fig. 8), a condition not found in other aplocheiloid fishes; 3) bony laminar ventral process close to the lateral Fig. 8. Diagrammatic representation of the head, dorsal view, of Kryptolebias caudomarginatus, UFRJ 6253, male, 48.5 mm SL. Drawn by the author Fig. 9. Fifth ceratobranchial, ventral view, of Kryptolebias caudomarginatus, UFRJ 6236, male, 38.5 mm SL. Drawn by the author 11 aqua vol. 11 no

14 Redescription of Kryptolebias ocellatus (Hensel) and K. caudomarginatus (Seegers) (Teleostei: Cyprinodontiformes: Rivulidae) edge of the fifth ceratobranchial (Fig. 9), not found elsewhere among aplocheiloid fishes. Acknowledgements I am grateful to C. P. Bove, B. B. Costa, T. Litz, J. P. Moreira, A. Moreira, J. Barata, J. L. Mattos, L. Villa Verde, F. Ottoni, and E. Mattos for help during collections. This study was also supported by CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico Ministério de Ciência e Tecnologia) and FAPERJ (Fundação de Amparo à Pesquisa do Estado do Rio de Janeiro). References Costa, W. J. E. M Comments on the proposed conservation of the specific name of Rivulus marmoratus Poey, 1880 (Osteichthyes, Cyprinodontiformes). Bulletin of Zoological Nomenclature, 51: Costa, W. J. E. M Pearl killifishes - the Cynolebiatinae: systematics and biogeography of the neotropical annual fish subfamily. TFH, Neptune City, 128 pp. Costa, W. J. E. M Phylogeny and classification of Rivulidae revisited: evolution of annualism and miniaturization in rivulid fishes (Cyprinodontiformes: Aplocheiloidei). Journal of Comparative Biology, 3: Costa, W. J. E. M The neotropical annual fish genus Cynolebias (Cyprinodontiformes: Rivulidae): phylogenetic relationships, taxonomic revision and biogeography. Ichthyological Exploration of Freshwaters, 12: Costa, W. J. E. M Family Rivulidae (South American annual fishes). In: R.E. Reis, Kullander, S.O. & Ferraris, C.J., Jr. (eds) Check list of the freshwater fishes of South and Central America, Edipucrs, Porto Alegre, pp Costa, W. J. E. M. 2004a. Relationships and redescription of Fundulus brasiliensis (Cyprinodontiformes: Rivulidae), with description of a new genus and notes on the classification of the Aplocheiloidei. Ichthyological Exploration of Freshwaters, 15: Costa, W. J. E. M. 2004b. Kryptolebias, a substitute name for Cryptolebias Costa, 2004 and Kryptolebiatinae, a substitute name for Cryptolebiatinae Costa, 2004 (Cyprinodontiformes: Rivulidae). Neotropical Ichthyology, 2: Costa, W. J. E. M. 2004c. A new killifish genus and species from the coastal plains of north-eastern Brazil (Teleostei: Cyprinodontiformes: Rivulidae). Zootaxa, 642: Davis, W. P., D. S., Taylor & B. J. Turner Field observations of the ecology and habits of mangrove rivulus (Rivulus marmoratus) in Belize and Florida (Teleostei: Cyprinodontiformes: Rivulidae). Ichthyological Exploration of Freshwaters, 1: Harrington, R. W Oviparous hermaphroditic fish with internal self-fertilization. Science, 134: Hensel, R Beiträge zur Kenntnis der Wirbelthiere Südbrasiliens. Archiv für Naturgeschichte Wiegman, 34: Hoedeman, J. J The frontal scalation pattern in some groups of toothcarps (Pisces, Cyprinodontiformes). Bulletin of Aquatic Biology, 1: Hrbek, T. & A. Larson The diapause in the killifish family Rivulidae (Atherinomorpha, Cyprinodontiformes): a molecular phylogenetic and biogeographic perspective. Evolution, 53: Huber, J. H Review of Rivulus: ecobiogeografia - relationships. Cybium, Société Française d Ichtyologie, Paris, pp. International Commission of Zoological Nomenclature Rivulus marmoratus Poey, 1880 (Osteichthyes, Cyprinodontiformes): given precedence over R. ocellatus Hensel, 1868, and a neotype designated for R. marmoratus. Bulletin of Zoological Nomenclature, 52: Lazara, K. J. & M. L. Smith 1990 Rivulus marmoratus Poey, 1880 (Osteichthyes, Cyprinodontiformes): proposed conservation of the specific name. Bulletin of Zoological Nomenclature, 47: Murphy, W. J., J. E. Thomerson & G. E. Collier Phylogeny of the neotropical killifish family Rivulidae (Cyprinodontiformes, Aplocheiloidei) inferred from mitochondrial DNA sequences. Molecular and Phylogenetic Evolution, 13: Notare, M Peixes raros do Brasil. Revista de Aquariofilia, 5: Poey, F Revisio piscium cubensium. Anales de la Sociedad Española de Historia Natural, 9: Seegers, L Zur Revision der Rivulus-Arten Südost-Brasiliens, mit einer Neubeschreibung von Rivulus luelingi n. sp. und Rivulus caudomarginatus n. sp. (Pisces: Cyprinodontidae: Rivulinae). Zoologische Beiträge, 28: Taylor, S Observations on the ecology of the killifish Rivulus marmoratus (Cyprinodontidae) in an infrequently flooded mangrove swamp. Northeast Gulf Science, 10: Taylor, S Adaptive specializations of the cyprinodont fish Rivulus marmoratus. Biological Sciences, 53: Taylor, W. R. & G. C. Van Dyke Revised procedures for staining and clearing small fishes and other vertebrates for bone and cartilage study. Cybium, 9: Turner, B. J., W. P. Davis & D. S. Taylor Abundant males in populations of a selfing hermaphroditic fish, Rivulus marmoratus, from Belize cays. Journal of Fish Biology, 40: Vermeulen, F. B. M. & T. Hrbek Kryptolebias sepia n. sp. (Actinopterygii: Cyprinodontiformes: Rivulidae), a new killifish from the Tapanahony River drainage in southeast Surinam. Zootaxa, 928: aqua vol. 11 no

15 aqua, Journal of Ichthyology and Aquatic Biology Centropyge abei, a new species of deep-dwelling angelfish (Pomacanthidae) from Sulawesi, Indonesia Gerald R. Allen 1, Forrest Young 2 and Patrick L. Colin 3 1) Western Australian Museum, Perth, Australia; mailing address: 1 Dreyer Road, Roleystone, WA 6111, Australia. 2) th Avenue Gulf, Marathon, Florida 33050, USA. 3) Coral Reef Research Foundation, Box 1765, Koror, PW 96940, Palau. Accepted: Keywords Taxonomy, marine fishes, Centropyge, new species, Pomacanthidae, Indonesia Abstract A new species of pomacanthid fish, Centropyge abei, is described from a single specimen, 90.8 mm SL, collected from a depth of 120 m during deep diving operations off Manado, Sulawesi in Indonesia during April It was also observed in similar habitat at Palau between m using a research submersible. The combination of morphological features that include large scales (43-45 in lateral row from upper opercle to caudal fin base), 3-4 preopercular spines, a relatively narrow supracleithrum with pronounced serrae on its posterior upper margin, serrate interoperculum, and a posterior margin of the preorbital bone that is attached and hidden by skin and scales serve to distinguish it from all known angelfishes, although it appears to be a modified Centropyge. Moreover, its distinctive colour pattern consisting of an extensive black area on the back and dorsal fin that grades to yellow on the side, white bar behind the head, and white caudal fin, is unlike that of any known species in the family. Zusammenfassung Beschrieben wird die neue Kaiserfischart Centropyge abei nach einem einzelnen Exemplar mit 90,8 mm SL, das in einer Tiefe von 120 Metern bei Tieftauchgängen vor Monado, Sulawesi, in Indonesien im April 2005 gefangen wurde. Weitere Exemplare konnten in einem ähnlichen Habitat bei Palau in einer Tiefe von 110 bis 155 Metern von einem Forschungstauchboot aus beobachtet werden. Große Schuppen (43-45 in einer seitlichen Reihe vom oberen Operculum bis zur Schwanzflossenbasis), 3-4 Präoperculum-Stacheln, ein relativ schmales Supracleithrum mit deutlichen Sägezähnchen am hinteren oberen Rand, ein gesägtes Interoperculum sowie ein Hinterrand des Präorbitalknochens, den anheftende Haut und Schuppen verdecken das sind Merkmale, die insgesamt diese neue Art von allen anderen Kaiserfischen unterscheiden, wobei sie schon als zu Centropyge gehörig erscheint. Auch die einzigartige Farbgebung unterscheidet sich von allen anderen Angehörigen der Familie: eine ausgedehnte schwarze Fläche am Rücken und auf der Rückenflosse, das an der Seite in einen Gelbton übergeht, ein weißer Streifen hinter dem Kopf und weiße Schwanzflosse. Résumé On décrit une nouvelle espèce de Pomacanthidé, Centropyge abei à partir d'un seul spécimen, d'une LS de 90,8 mm, collecté à une profondeur de 120 m au cours de plongées profondes au large de Manado, Sulawesi, en Indonésie, au mois d'avril Cette espèce a aussi été observée, dans un habitat similaire, à Palau, entre 110 et 155 m, à l'aide d'un sous-marin de recherche. Une combinaison de caractéristiques morphologiques: de larges écailles (43-45 sur le rang horizontal du haut de l'opercule à la base de la caudale), 3-4 rayons durs préoperculaires, un supercleithrum relativement étroit avec des dentelures prononcées sur la marge supérieure postérieure, un interopercule dentelé et une marge postérieure de l'os préorbital qui est fixée et cachée par la peau et les écailles, aident à le distinguer de tous les poissons-anges, même si elle ressemble à un Centropyge altéré. En outre, son patron de coloration caractéristique: une large zone noire sur le dos et la dorsale qui devient jaune sur le flanc, une barre blanche derrière la tête et une caudale blanche, ne ressemble à aucun de ceux d'espèces connues dans la famille. Sommario Una nuova specie di pomacantide, Centropyge abei, è descritta sulla base di un singolo esemplare di 90.8 mm SL, raccolto a 120 m di profondità durante immersioni profonde al largo di Manado, Sulawesi in Indonesia nell Aprile E stato anche osservato in un habitat simile a Palau a profondità di m utilizzando un sottomarino da ricerca. La combinazione di tratti morfologici che includono larghe scaglie (43-45 in linea laterale dall opercolo superiore alla base della pinna caudale), 3-4 spine preopercolari, un supracleitrum relativamente stretto con pronunciate dentellature sul suo margine superiore, interopercolo seghettato, e margine posteriore dell osso preopercolare attaccato e 13 aqua vol. 11 no

16 Centropyge abei, a new species of deep-dwelling angelfish (Pomacanthidae) from Sulawesi, Indonesia nascosto dalla pelle e dalle scaglie, servono a distinguerlo da tutti glia altri pesci angelo, sebbene appaia un Centropyge modificato. Inoltre, la distinta colorazione consistente in un estesa area nera sul dorso e sulla pinna dorsale che degrada nel giallo sui fianchi, una barra bianca dietro la testa e una pinna caudale bianca risulta diversa da tutte le specie note della famiglia. Introduction Angelfishes of the family Pomacanthidae are among the most conspicuous inhabitants of coral reefs, occurring in both shallow and relatively deep water. Allen et al. (1998) reviewed the family, recognizing 83 species worldwide, including 70 from the Indo-Pacific region. A more recent (2003) treatment of the group by Debelius et al. recognized 75 Indo-Pacific species, including 28 Indo-Pacific members of the genus Centropyge Kemp, The present paper describes a new species of deepdwelling angelfish that is provisionally assigned to the genus Centropyge. It was first sighted during submersible dives along the outer slope of Palau in 2001 by the third author. A single specimen, described herein, was subsequently collected during a series of deep rebreather scuba dives by the second author off Manado, Sulawesi in Indonesia during April Although the primary goal of this expedition, to locate and film a recently described coelacanth, was unsuccessful, several valuable collections of small reef fishes were obtained. Materials and Methods The specimen of the new angelfish was obtained with the aid of mixed gas, re-breather scuba equipment allowing dives to working depths of approximately 150 m. A mild anaesthetic, quinaldine sulphate, was used to partially narcotise fish specimens prior to capture with a hand net. Standard length (SL) was measured from the front of the upper lip to the base of the caudal fin (end of hypural plate). Total length (TL) was measured from the front of 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. Measurements 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 known specimen - is deposited at Pusat Penelitian dan Pengembangan Oseanologi, Jakarta, Indonesia (NCIP). Centropyge abei n. sp. (Figs 1-2) Holotype: NCIP 6305, 90.8 mm SL, south-east side of Manado Tua (approximately S, E), Sulawesi, Indonesia, 120 m, collected with quinaldine and hand net by F. Young, 27 April Diagnosis A species of the pomacanthid genus Centropyge with the following combination of characters: dorsal rays XIII,17; anal rays III,18; pectoral rays 16; scales relatively large, in lateral row from upper end of gill opening to base of caudal fin; lateral line with 37 scales, terminating below end of dorsal fin, but re-appearing on 7 scales of caudal peduncle; gill rakers on first arch ; maximum body depth 1.6 in SL; preopercular spines 3 or 4; length of primary preopercular spine 2.0 in head length; teeth tricuspid, about in outer row in each jaw; exposed margins of preorbital, interopercle, preopercle, subopercle, supracleithrum, and posttemporal bones serrate; posterior margin of preorbital bone attached and hidden by skin and scales; live coloration yellow with black upper third of back, dorsal fin, and upper part of head. A broad white bar immediately anterior to dorsal fin origin, extending to at least level of upper opercular margin. Caudal fin and peduncle white. Description Dorsal rays XIII,17 (last divided to base); anal rays III,18 (last divided to base); pectoral rays 16 (upper two and lowermost unbranched); pelvic rays I,5; transverse scale rows from upper end of gill opening to base of caudal fin 43-45; scale rows above lateral line to origin of dorsal fin 9; scale rows below lateral line to origin of anal fin about 20; lateral line in two sections, the anterior part extending from upper edge of gill cover to below level of posteriormost dorsal fin ray containing 37 scales and the posterior part containing 7 scales along middle Fig. 1. Freshly collected specimens of Centropyge abei n. sp., holotype, 90.8 mm SL, Manado Tua, Sulawesi, Indonesia. Photo by Kotaro Yoshimura. aqua vol. 11 no

17 Gerald R. Allen, Forrest Young and Patrick L. Colin of caudal peduncle; transverse scale rows on opercle 7; gill rakers on first branchial arch , total rakers 17, the rakers short and stubby, much shorter than gill filaments; vertebrae Body ovate, the depth 1.6 (61.3 % of SL) in SL, and compressed, the maximum width 2.6 (21.4 % of SL) in depth; head length 3.2 (30.9 % of SL) in SL; dorsal profile of forehead relatively steep and straight, forming an angle of about 45 degrees to the horizontal; snout 2.6 (12.1 % of SL) in head length, diameter of orbit 3.1 (10.0 % of SL) in head length; interorbital space slightly convex, the least bony width 3.5 (8.8 % of SL) in head length; caudal peduncle much deeper than long, the least depth 2.2 (14.3 % of SL) in head length; length of caudal peduncle 6.0 (5.2 % of SL), in head length. Mouth relatively small, terminal, the gape forming an angle of about 30 degrees to the horizontal, the maxilla reaching a vertical at about front of anterior nostril. Upper and lower lips about equal in size, broadly scaled except anterior edges, their maximum width contained 2.8 in diameter of orbit. Teeth slender, elongate (the longest 4.6 in orbit), close-set, flexible, tricuspid (large central cusp notably longer than smaller lateral ones), in 5-6 rows in each jaw, about in outer row of each jaw. No teeth on roof of mouth. Tongue short and broadly rounded. Nostrils anterior to centre of eye, the posterior opening relatively large and ovate with thin, slightly raised rims; anterior nasal opening about one-fourth size of posterior one, in a membranous tube with a posterodorsal flap; Most head pores inconspicuous (due to dense covering of tiny scales) except for prominent pore just anterior to anterior nasal opening. Gill membranes narrowly attached to isthmus. Longest gill filament on first arch contained 1.9 times in orbit. Gill rakers short, the longest 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

18 Centropyge abei, a new species of deep-dwelling angelfish (Pomacanthidae) from Sulawesi, Indonesia Upper edge of opercle obtusely rounded, without a spine. A prominent large spine at corner of preopercle, longer than orbit, the spine length (measured along lower edge) contained 2.0 (15.3 % of SL) times in head length; a series of 2 (right side) or 3 smaller preopercle spines immediately below and anterior to primary spine, the largest (posteriormost) one-fourth (right side) to onehalf length of primary spine; margin of preopercle with serrae; anterior and ventral margins of preorbital with 5-9 serrae, the posterior margin attached and hidden by skin and scales; lower exposed margin of interopercle with 2-5 serrae; margin of subopercle with serrae; exposed posterior margin of supracleithrum relatively narrow and covered with scales, its upper posterior edge with serrae; posterior edge of posttemporal with 4-7 serrae. Scales of body more or less arranged in regular transverse rows and coarsely ctenoid (up to 25 ctenii on exposed posterior margins); auxiliary scales mainly confined to lateral line and anteriormost portion of body; head fully scaled except anterior edges of lips; dorsal and anal fins scaled nearly to margins except anteriorly in spinous portion where fin membranes are deeply incised; caudal fin scaled nearly to posterior margin; pectoral and pelvic fins densely scaled at base, with tiny scales extending on surface of rays (but not on membranes) nearly to posterior margin. Lateral line relatively inconspicuous, its presence indicated by smaller scales (including tiny auxiliary scales), gently arching across back, originating at upper corner of gill opening and terminating below end of soft dorsal fin; additionally, 7 tubed scales midlaterally on caudal peduncle. Caudal fin rounded, its length 1.2 (25.3 % of SL) in head length. Origin of dorsal fin slightly anterior to a vertical at upper end of gill opening. Dorsal spines progressively longer to last spine; first dorsal spine 2.7 (11.5 % of SL), second dorsal spine 2.1 (14.5 % of SL), third dorsal spine 1.9 (16.0 % of SL), all in head length; membranes between first eight dorsal spines and three anal spines deeply incised; posterior margin of soft portions of dorsal and anal fins angular, the longest dorsal and anal rays reaching as far posterior as level of middle of caudal fin, their length 1.5 (21.0 % of SL) and 1.4 (22.8 % of SL) respectively in head length; origin of anal 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

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

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

21 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 , USA Accepted: Keywords Taxonomy, Gobiidae, Amblyeleotris, new species, New Britain Abstract The gobiid fish Amblyeleotris neumanni is described as a new species from three specimens, mm SL, collected in 26 m off Lolobau Island, New Britain, Papua New Guinea. It was observed in symbiotic association with the snapping shrimps Alpheus rapacida and A. sp., with which it shares a burrow. It is distinctive in having 12 or 13 dorsal soft rays, 13 anal soft rays, 19 pectoral rays, scales in longitudinal series, males with filamentous third to fifth spines, lanceolate caudal fin, pelvic fins fully joined, pelvic frenum, and in colour: four broad dark orangish brown bars on body with irregular dark markings between; ocellated orange spots on spines and rays of dorsal fins; basal part of anal fin with adjacent narrow longitudinal bands of black, orange, blue, and blue-green; pelvic fins pale blue or green. Zusammenfassung Beschrieben wird als neue Art die Meergrundel Amblyeleotris neumanni nach drei Exemplaren mit 44,2-53,9 mm SL, die in 26 Metern Tiefe vor den Lolobau- Inseln, New Britain, Papua-Neuguinea gefangen wurden. Sie wurden in Symbiose mit dem Knallkrebs Alpheus rapacida und einer weiteren Art beobachtet, mit denen sie die Höhle teilten. Ihre Unterscheidungsmerkmale sind 12 oder 13 weiche Rückenflossenstrahlen, 13 weiche Analflossenstrahlen, 19 Brustflossenstrahlen, Schuppen in Längsreihen, bei den Männchen fädige Stacheln an 3. und 5. Stelle, eine lanzettförmige Schwanzflosse, vollständig vereinte Bauchflossen, ein Beckenband und die Farbgebung: vier breite, dunkle orangebraune Bänder auf dem Rumpf mit unregelmäßigen dunklen Markierungen dazwischen; orangefarbene Augenflecken auf Stacheln und Flossenstrahlen der Rückenflossen; am basalen Teil der Analflosse schmale Längsstreifen in Schwarz, Orange, Blau und Blaugrün; Bauchflossen blassblau oder -grün. Résumé Le gobie Amblyeleotris neumanni est décrit en tant qu'espèce nouvelle sur base de trois spécimens, d'une LS de 44,2 à 53,9 mm, et a été collecté, à 26 m, au large de l'île Lolobau, Nouvelle Angleterre, Papouasi Nouvelle Guinée. Il a été observé en association symbiotique avec les crevettes Alpheus rapacida et A. sp., avec lesquelles il partage un terrier. Il se caractérise par la présence de 12 ou 13 rayons mous à l'anale, 19 rayons pectoraux, écailles en rangée longitudinale, les mâles, avec les 3e et 5e rayons filamenteux, une caudale lancéolée, des nageoires pelviennes complètement soudées, un frenum pelvien, et par la coloration: quatre larges barres brun orange sur le corps entrecoupées de taches foncées irrégulières, des ocelles orange sur les rayons durs et des rayons mous des dorsales, la base de l'anale marquée longitudinalement de lignes étroites adjacentes, noire, orange, bleue et bleuvert; pelviennes bleu pâle ou vertes. Sommario Il gobide Amblyeleotris neumanni è descritto come specie nuova sulla base di tre esemplari di mm SL, raccolti a 26 m al largo dell isola di Lolobau, Nuova Britannia, Papua Nuova Guinea. E stato osservato in associazioni simbiotiche con il gamberetto Alpheus rapacida e A. sp., con cui condivide la tana. Si distingue epr avere 12 o 13 raggi molli dorsali, 13 raggi molli anali, raggi 19 pettoral, scaglie in serie longitudinale, i maschi con terza, quarta e quinta spina filamentosa, pinna caudale lanceolata, pinne pelvicche completamente unite, frenum pelvico presente e la colorazione: quattro ampie barre arancio-marrone scuro sul corpo con macchie irregolari inframmezzate; macchie arancio ocellate sulle spine e i raggi della dorsale; base della pinna anale con strette bande ravvicinate longitudinali di colore nero, arancio, blu e grigio-blu; pinne pelviche blu pallido o verde. Introduction Species of 13 genera of gobiid fishes live in symbiotic association with snapping shrimps of the genus Alpheus. The shrimp builds and maintain a burrow in sedimentary substrata. The goby, with its superior vision and cephalic sensory system, serves as the sentinel at the burrow entrance, quickly taking refuge in the burrow, if needed. The most speciose of the 13 genera is the Indo-Pacific genus Amblyeleotris, which includes some of the most colourful fishes of our planet. It was described by Bleeker (1874), designating Eleotris periophthalmus Bleeker as the type 19 aqua vol. 11 no

22 Amblyeleotris neumanni, a new species of shrimp goby from New Britain species. In a paper describing five new species of Amblyeleotris from islands of Oceania, Randall (2005) provided a diagnosis of the genus and a brief historical resume of the species. A new species from the island of New Britain, Papua New Guinea is described in the present paper, bringing the total number of species in the genus to 34. The opportunity to collect and photograph fishes in New Britain was provided by a cruise on the dive vessel Pelagian, which commenced in July, 2002 at Kimbe Bay on the north coast and sailed east to Rabaul. About half way to Rabaul we stopped to dive at Lolobau Island, an active volcano 6.5 by 10 km in size and 832 m high. We were joined on the dive by Mike Neumann who had financed the cruise. We spotted a new species of Amblyeleotris on a bottom of dark brown sand and rubble at a depth of 26 m. Underwater photographs were taken, and three specimens were collected with a small multiprong Hawaiian sling spear. Materials and Methods The holotype and one paratype of the new species are deposited in the Bishop Museum, Honolulu (BPBM), and one paratype in the U.S. National Museum of Natural History, Washington, D.C. (USNM). The length of specimens is given as standard length (SL), measured from the median anterior end of the upper lip to the base of the caudal fin (posterior end of the hypural plate); body depth from the origin of pelvic fins, and body width at the origin of the pectoral fins; head length (HL) from the upper lip to the posterior end of the opercular membrane, and head width over the posterior margin of the preopercle; orbit diameter is the greatest fleshy diameter, and interorbital width the least bony width; snout length is measured from the median anterior point of the upper lip to the nearest fleshy edge of the orbit; upper jaw length from the same anterior point to the posterior end of the maxilla; caudal peduncle depth is the least depth, and caudal peduncle length the horizontal distance between verticals at the rear base of the anal fin and the caudal fin base; lengths of spines and rays are measured to their extreme bases; caudal fin and pectoral fin lengths are the length of the longest ray; pelvic fin length is measured from the base of the pelvic spine to the tip of the longest soft ray. Morphometric data are presented in Table I as percentages of the standard length. Proportional measurements in the text are rounded to the nearest The count of scales in longitudinal series is made from above the upper end of the gill opening to the base of the caudal fin; scales in transverse series are counted from the origin of the anal fin obliquely upward to the base of the first dorsal fin; the count of gill rakers is made on the first gill arch, those on the upper limb given first. Meristic and morphometric data in parentheses refer to paratypes. Amblyeleotris neumanni n. sp. (Figs. 1-4; Table I) Holotype: BPBM 39048, 54.9 mm, Papua New Guinea, New Britain, Lolobau Island, east end off red cliff, S, E, dark silty sand and rubble, 26 m, collected with spear by J. E. Randall, 3 August Paratypes: BPBM 39049, 44.2 mm and USNM , 46.3 mm, same data as holotype. Diagnosis Dorsal rays VI+I,12-13; anal rays I,13; pectoral rays 19; longitudinal scale series ; no median predorsal scales; scales on side of nape extending forward to above middle of opercle; body depth in SL; head length in SL; snout length 1.75 in HL; gill opening extending forward to above middle of peopercle; third to fifth dorsal spines of male prolonged as filaments; caudal fin lanceolate, in SL; pelvic fins fully joined medially, reaching origin of anal fin; pelvic frenum present; four orangish brown bars on body with irregular dark markings between; rays of dorsal fins with ocellated orange spots, the soft rays tipped with orange; basal part of anal fin with narrow adjacent bands of black, orange, blue, and blue-green; pelvic fins pale blue or green. Description Dorsal rays VI+I,13 (12-13); anal rays I,13; all dorsal and anal soft rays branched, the last to base; pectoral rays 19, the uppermost and lower 1 or 2 unbranched; pelvic rays I,5, all soft rays branched, the medial rays fully united by membrane; pelvic frenum present; segmented caudal rays 17, 14 (13-14) branched; upper and lower unsegmented caudal rays 7; longitudinal scale series 98 (99-100); transverse scale rows 31 (32-33); circumpeduncular scales about 25; gill rakers 4+13 ( ); pseudobranch with 8 fleshy lobes; branchiostegal rays 5; vertebrae 25. Body depth 5.8 ( ) in SL; body width 1.75 ( ) in body depth; HL 3.5 ( ) in SL; head width 2.5 ( ) in HL; snout length 1.75 (1.75) in HL; dorsal profile of snout moderately steep, forming an angle of about 40 to horizontal axis of head and body; orbit diameter 3.95 ( ) in HL; upper edge of eye extending above dorsal profile of head; interorbital space extremely narrow, the least width 36 (41) in HL; caudal peduncle depth 3.0 ( ) in HL; caudal peduncle length 1.75 ( ) in HL. Mouth oblique, forming an angle of about 35 to horizontal axis of head and body, with the lower jaw slightly projecting; maxilla reaching to or slightly pos- aqua vol. 11 no

23 John E. Randall and John L. Earle terior to a vertical through centre of eye, the upper jaw length 2.6 ( ) in HL; front of upper jaw with two well-spaced, incurved canine teeth, the symphyseal gap in holotype nearly equal to pupil diameter, the length of the largest and more medial of the canines two-thirds pupil diameter in length; side of upper jaw with a row of 18 slender incurved canine teeth in holotype, progressively shorter posteriorly; an inner band of small, strongly incurved villiform teeth in three or four rows at front of jaws, narrowing to a single row posteriorly; front of lower jaw with three pairs of incurved slender canine teeth smaller than those of upper jaw and an inner band of incurved villiform teeth in at most three rows; side of lower jaw nearly halfway from symphysis with three large close-set retrorse canine teeth, the largest as long as largest canine at front of upper jaw; remainder of lower jaw with a single row of close-spaced slender teeth smaller than those of upper jaw; no teeth on vomer or palatines; tongue broadly rounded; inner edge of lips papillose. Gill opening extending forward to below middle of preopercle (about one-half eye diameter posterior to eye). Gill rakers short, the longest about half length of longest gill filaments. Posterior nostril anterior to lower third of eye just before fleshy edge of orbit, the aperture small, only slightly larger than sensory pores of head, with a very low rim; anterior nostril ventroanterior to posterior nostril, at end of a short membranous tube, nearly reaching edge of snout when depressed forward. Pores of cephalic sensory system and sensory papillae on cheek and opercle as shown in illustration of Amblyeleotris japonica by Akihito in Masuda et al. (1984: 255, fig. 95). Scales small for the genus, progressively smaller anteriorly, extending on side of nape to above middle of opercle, but none in median predorsal zone or prepectoral area (if embedded scales are present in these two apparently naked areas, none could be dislodged); scales ctenoid posteriorly on body, becoming cycloid anterior to origin of second dorsal fin; small cycloid scales present on chest; no scales on fins except basal fourth to fifth of caudal fin. Origin of first dorsal fin over inner base of pelvic fins, the predorsal length 3.4 (3.3) in SL; first dorsal spine 1.8 ( ) in HL; fourth and fifth dorsal spines longest and subequal, 1.2 (1.2 in male paratype and 1.6 in female paratype) in HL; third to fifth dorsal spines prolonged as short filaments in males; spine of second dorsal fin 2.2 ( l) in HL; penultimate and preceding dorsal soft rays subequal and longest, 1.55 ( ) in HL; origin of anal fin a little anterior to base of second dorsal soft ray, preanal length 1.8 (1.8) in SL; anal spine 3.45 ( ) in HL; penultimate and preceding anal soft rays longest, 1.5 (1.6) in HL; caudal fin lanceolate, 2.6 ( ) in SL; pectoral rays pointed, the middle rays longest, 1.15 ( ) in HL; prepelvic length 3.4 ( ) in SL; pelvic spine 2.65 ( ) in HL; fifth pelvic soft ray longest, extending to origin of anal fin, 3.7 ( ) in SL; pelvic frenum well developed, the membrane reaching about two-thirds length of pelvic spine. Colour of male holotype in life: shown in Fig. 1. A smaller male is illustrated in Fig. 2, and females in Figs. 3 and 4. Noteworthy colour features are the four dark orangish brown bars on the body with irregular 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

24 Amblyeleotris neumanni, a new species of shrimp goby from New Britain Table I. Proportional measurements of type specimens of Amblyeleotrus neumanni n. sp. as percentages of the standard length. Holotype Paratypes BPBM BPBM USNM Sex male female male Standard length (mm) Body depth Body width Head length Head width Snout length Orbit diameter Interorbital width Upper jaw length Caudal peduncle depth Caudal peduncle length Predorsal length Preanal length Prepelvic length Base of dorsal fins First dorsal spine Longest dorsal spine Spine of second dorsal fin Longest dorsal ray Base of anal fin Anal spine Longest anal ray broken Caudal fin length Pectoral fin length Pelvic spine length Pelvic fin length dark markings in the bluish to whitish interspaces, the dark brown, blue, and orange markings on the head, the ocellated orange spots on the dorsal spines (the spines of females tipped with orange), the second dorsal fin with an orange spot just posterior to ray tips, the basal part of anal with adjacent longitudinal bands of black, orange, blue, and blue-green, the orange upper margin of the caudal fin, and the pale blue pelvic fins. Colour of holotype in alcohol: pale tan, becoming still paler ventrally, with four broad brown bars narrower than pale interspaces, the first on nape narrowing ventrally onto opercle, and the last on caudal peduncle; pale interspaces with irregular narrow vertical dark bands dorsally on body, some commencing on back in a distinct brown spot; interorbital space and upper edge of eyes dark brown, the pigment following sensory canal behind upper part of eye and continuing horizontally as a band to upper pore of preopercular sensory canal; front of snout and medial part of lips and chin brown; a faint brown bar from below centre of eye to behind posterior end of upper jaw; ventral margin of gill membranes broadly dark brown; dorsal fins pale yellowish, the first dorsal dusky with a pale spots (fin damaged on holotype; spotting more evident on fin of male paratype); caudal fin pale, a little dusky on scaled basal part, with a dark brown streak on outer part of membrane between tenth and eleventh branched rays; anal fin transparent with a narrow, very dark brown stripe in lower middle part of fin, bordered above by a light brown band that is bisected by a darker brown line; pectoral fins pale, Fig. 2. Amblyeleotris neumanni n. sp., male, with Alpheus sp., Lolobau Island, New Britain. Photo by J. E. Randall. aqua vol. 11 no

25 John E. Randall and John L. Earle the rays edged with light brown, with a dusky spot on upper base; pelvic fins with brown membranes that are darker medially. Etymology We are pleased to name this colourful goby in honour of Mike Neumann, fellow diver, underwater photographer and good friend. Remarks This species was observed and collected only at one site in 26 m at Lolobau Island, New Britain. Figures 2 and 3 show its symbiotic partner Alpheus sp., probably an undescribed species related to A. brevirostris (Arthur Anker, pers. comm.). A second snapping 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

26 Amblyeleotris neumanni, a new species of shrimp goby from New Britain shrimp, A. rapacida, is illustrated in Fig. 4. Amblyeleotris neumanni is most similar to A. masuii Aonuma and Yoshino, described from Okinawa in It was illustrated in colour by Hayashi and Shiratori (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, % HL, compared to % 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 measuring 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 reddish brown, and the second dorsal rays tipped in dark reddish brown), the dark brown band behind the middle of the eye of neumanni, and the four different coloured bands on the base of the anal fin of neumanni (only a single orange line in masuii). Japan. 534 pp. Heibonsha, Ltd., Tokyo (in Japanese). Acknowledgements We thank Mike Neumann for his assistance in collecting 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 Two new species of the genus Amblyeleotris (Pisces: Gobiidae) from the Ryukyu Islands, Japan. Ichthyological Research, 43 (2): Bleeker, P Notice sur les genre Amblyeleotris, Valenciennesia et Brachyeleotris, Verslagen en Mededeelingen der Koninklijke Akademie van Wetenschappen, 8: Hayashi, M. & T. Shiratori, Gobies of Japanese Waters. 223 pp. TBS Buritanica, Tokyo (in Japanese). Masuda, H., Amaoka, K., Araga, C., Uyeno, T. & T. Yoshino (Eds.) The Fishes of the Japanese Archipelago. Vol. 1 (text xxii pp) and vol. 2 (plates). Tokai University Press, Tokyo. Randall, J. E Five new shrimp gobies of the genus Amblyeleotris from islands of Oceania. aqua, Journal of Ichthyology and Aquatic Biology, 8 (2): Senou, H., Suzuki, T., Shibukawa, K. & K. Yano A Photographic Guide to the Gobioid Fishes of aqua vol. 11 no

27 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. wivantso@rna.bio.mq.edu.au Accepted: Keywords Isonidae, phylogenetic analysis, anatomy, osteology Abstract The musculoskeletal anatomy of Iso rhothophilus is described. A phylogenetic analysis of the genus, using eleven anatomical characters, indicates that the systematic hierarchy is (Iso flosmaris (I. nesiotes (I. rhothophilus (I. hawaiiensis, I. natalensis)))). Isonidae Rosen, 1964 is redefined and, on the basis of twenty characters (autapomorphic within Atheriniformes) shown to be distinct from Notocheiridae Schultz, Zusammenfassung Beschrieben wird die Anatomie des Muskelskeletts von Iso rhothophilus. Nach einer phylogenetischen Analyse der Gattung anhand von elf anatomischen Merkmalen muss man auf die folgende systematische Hierarchie schließen: (Iso flosmaris (I. nesiotes (I. rhothophilus (I. hawaiiensis, I. natalensis)))). Die Isonidae Rosen, 1964, werden neu definiert und ihre Unterscheidung zu den Notocheiridae Schultz, 1950, auf der Grundlage von 20 Merkmalen (autapomorphisch innerhalb der Atheriniformes) festgestellt. Résumé On décrit l'anatomie musculo-squelettique d'iso rhothophilus. Une analyse phylogénétique du genre, à l'aide de onze caractéristiques anatomiques, montre que la hiérarchie systématique est (Iso flosmaris (I. nesiotes (I. rhothophilus (I. hawaiiensis, I. natalensis)))). Isonidae Rosen, 1964 est redéfini et, à l'aide de vingt caractéristiques (autapomorphiques pour les Atheriniformes), on montre sa différence par rapport à Notocheiridae Schultz, Sommario L anatomia muscoloscheletrica di Iso rhothophilus è descritta in dettaglio. Un analisi filogenetica del genere, eseguita con undici caratteri anatomici, indica che la gerarchia sistematica è (Iso flosmaris (I. nesiotes (I. rhothophilus (I. hawaiiensis, I. natalensis)))). La famiglia Isonidae Rosen, 1964 viene ridefinita e, sulla base di venti caratteri (autapomorfici entro gli Ateriniformes) si dimostra distinta dai Notocheiridae Schultz, Introduction Isonids are small fishes that have been infrequently collected in surf, usually near rocks, at the margins of the Pacific, Indian and Atlantic Oceans. The history of the genus Iso and the designation of the family Isonidae are fraught with confusion. Ogilby (1895) described the first species of the flower of the wave (as it was subsequently commonly called) as Tropidostethus rhothopilus from Maroubra Bay, NSW, Australia. In 1901, Jordan and Starks named another species as Iso flosmaris from Japan. A few years later Waite (1904) indicated that Iso was indistinct from Tropidostethus, then, in 1919, Jordan and Hubbs pointed out that Tropidostethus was preoccupied by a genus of insects thus determining the validity of the name Iso. Other species ascribed to the genus were I. natalensis Regan, 1919, I. flosindicus Herre, 1944, I. hawaiiensis Gosline (1952) and I. nesiotes Saeed, Ivantsoff & Crowley, A description of Notocheirus hubbsi by Clark in 1937 was noted by Hubbs (1944) as a species which was Iso-like but according to him was likely to be an offshoot of Atherinopsinae. Further confusion arose with the description of a subfamily of Tropidostethinae by Schultz in According to him this subfamily included Notocheirus, Tropidostethus and Iso, presumably considering that the type genus of the subfamily was Tropidostethus. In 1950 Schultz appears to have realised that Tropidostethus was preoccupied, having substituted the name with Tropidosthetops, at the same time changing the subfamilial name to Notocheirinae, and stating that the new subfamily was based on the genus Notocheirus Clark. In a review of Iso Jordan and Starks (Said, 1983), five species were considered valid, and the genus was considered distinct from Notocheirus Clark. Said (1983) noted that Isonidae Rosen, 1964 was erected on the basis of examination of Notocheirus hubbsi Clark, a monotypic genus previously assigned type status for Notocheirinae Schultz, In a review of the systematic position of Isonidae 25 aqua vol. 11 no

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

29 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

30 Descriptive anatomy of Iso rhothophilus (Ogilby), with a phylogenetic analysis of Iso and a redefinition of Isonidae (Atheriniformes) region. Larger specimens to 60 mm SL. Musculoskeletal anatomy Cranium: Rostrum elongate (Fig. 3). Ethmoid cartilage partially exposed laterally to form facet for articulation with palatine. Lateral ethmoid with cartilaginous condyle for articulation with lacrimal, supported dorsally and ventrally by osseous laminae. Lateral ethmoid with cartilaginous process contacting palatoquadrate cartilage, ventral to condyle for lacrimal. Lateral ethmoid closely contacting contralateral structure, not forming articulation with parasphenoid. Lateral ethmoid medial border with notch for olfactory nerve. Rostral cartilage bound to premaxilla dorsal process. Dorsal mesethmoid absent. Vomer with bilateral max- 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

31 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: parasphenoid; pf: palatine facet; pr: prootic; ps: pterosphenoid; vo: vomer. illary condyles (Fig. 4). Parasphenoid enlarged posteriorly, embedded in basioccipital. Parasphenoid anterior margin planar, interposed between ethmoid cartilage and vomer. Parasphenoid ventral ridge compressed, blade-like, extending posteriorly. Nasal irregular to triangular, separated from contralateral structure. Nasal cavity extending medially, separated from contralateral structure by thin ventromedian 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 ethmoid. Interorbital portion of frontal separated from supraorbital laminae by deep sulcus forming supraorbital 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). Posttemporal fossa formed by pterotic and sphenotic, and in part by epiotic and parietal, bounded laterally by osseous temporal canal. Temporal canal fused to pterotic, extending anteriorly to form junctions with supraorbital and infraorbital canals. Foramen situated at anterior temporal canal, superficial to sphenotic, forming nexus between dilator operculi fossa and canal. Dilator operculi fossa giving origin, in part, to levator operculi muscle. Anterior wall of fossa formed by sphenotic postorbital process. Sphenotic and pterotic forming facets articulating with hyomandibula. Subtemporal fossa medial and deep to posttemporal fossa, giving origin to branchial A B Fig. 5. Iso rhothophilus, MU I-122, 50.1 mm SL. Neurocranium in A: dorsal aspect; B: caudal aspect. Abbreviations: bo: basioccipital; ec: ethmoid cartilage; eo: exoccipital; eoc: exoccipital crest; ep: epiotic; fm: foramen magnum; fr: frontal; hf: hyomandibular face; lc: lacrimal; le: lateral ethmoid; me: mesethmoid; mo: myodome opening; na: nasal; oc: occipital condyle; of: olfactory nerve foramen; pa: parasphenoid; pc: epiotic crest; pe: pterotic; pf: palatine facet; pi: parietal; pt: posttemporal fossa; sc: supraorbital canal; so: supraoccipital; sp: sphenotic; tc: temporal canal; vf: vagus foramen; vo: vomer. Scale bar: 1mm. 29 aqua vol. 11 no

32 Descriptive anatomy of Iso rhothophilus (Ogilby), with a phylogenetic analysis of Iso and a redefinition of Isonidae (Atheriniformes) A B 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. levator musculature. Pterosphenoid separated from contralateral structure, lateral margin contacting frontal. Pterosphenoid, and ventral ridge originating on frontal, supporting membrane enclosing anterior lobe of brain. Membrane attached posteriorly to prootic and basisphenoid. Basisphenoid bearing bilateral posterolateral rami contacting prootic. Orbits separated medially by septum attached to parasphenoid. Ventromedian projection from prootics extending between bilateral rami of basisphenoid. Prootic jugular canal deep to osseous trabeculum. Posterior jugular opening confluent with emanation of hyomandibular nerve. Otic bulla small, extending ventrolaterally to parasphenoid. Pterotic bearing projection superficial to exoccipital. Occipito-vertebral articulation formed by spherical basioccipital condyle and paired small, separate exoccipital condyles. Jaws: Premaxilla with moderate symphysis, alveolar ramus slightly smaller than maxilla (Fig. 6). Premaxilla postmaxillary processes supporting skin fold between premaxilla and maxilla. Premaxilla teeth forming single row. Maxilla narrow, positioned lateral to premaxilla. Maxilla proximal angle narrow, distal angle broad. Maxilla bearing large condyle for articulation with vomer, biconcave submaxillary meniscus interposed between maxilla and vomer. Maxilla dorsal and internal processes formed about premaxilla dorsal process. Maxilla joined anteriorly to contralateral structure ligamentously. Premaxilla process providing attachment for articularmaxillary ligament (Fig. 7), originating from articular lateral face. Rostral cartilage supporting premaxilla dorsal process. No ligaments between maxilla and palatine. Ethmomaxillary ligament originating on mesethmoid, inserting on maxilla dorsal process. Labial ligament between anterolateral face of dentary and distal angles of maxilla and premaxilla. Coronomaxillary ligament absent. Adductor mandibulae muscle inserting in triangular recess dorsal to Meckelian cartilage between dentary and articular. Meckelian cartilage elongate (Fig 8), joining dentary and articular medially. Coronomeckelian bone, on dorsal surface of Meckelian cartilage adjacent to articular, acting as insertion for tendon of adductor mandibulae muscle. Articular with retrorse condyle on posterodorsal facet serving as insertion for ligament from quadrate. Retroarticular bone small, at angle of lower jaw, overlapping medially with articular. Suspensorium: Comprising palatoquadrate arch, ectopterygoid, mesopterygoid, ectopterygoid, symplectic, hyomandibula and preopercle (Fig 9). Two articulations formed with cranium, bilaterally: anterior articulation with palatine, posterior articulation with aqua vol. 11 no

33 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: ethmomaxillary 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. hyomandibula. Palatine and quadrate joined about interposed cartilage, cartilage also contacting ethmoid. Palatine cartilage progressively ossifying. Palatine maxillary process comprising cylindrical bone with cartilaginous core. Palatine bearing posteroventral process overlapping ectopterygoid. Mesopterygoid expansive, thin, forming ventromedian limit of orbit. Metapterygoid and quadrate joined about narrow interposed cartilage. Quadrate and symplectic with elongate articulation, both bones with broad ventral facets articulating with preopercle. Metapterygoid not articulating with cranium. Hyomandibula with ventral projection joined to preopercle ventral ramus. Hyomandibula articulating dorsally with cranium, hyomandibula posterior condyle articulating with opercle. Opercular series: Comprising opercle, interopercle and subopercle. Opercle large, elongate, bearing glenoid fossa articulating with hyomandibula. Dilator operculi muscle inserting on medial face of triangular dilator process of sphenotic and pterotic. Levator operculi muscle inserting on dorsal and medial surfaces of opercle. Interopercle wide, bearing dorsal process. Interopercle and branchiostegal rays covering gills ventrally. Interopercle joined to lower jaw ligamentously. Interopercle joined to ceratohyal by short ligament. Subopercle elongate, forming portion of gill cover. A B Fig. 8. Iso rhothophilus, MU I-122, 52 mm SL. Scanning electron micrograph of A: right premaxilla in lateral aspect; B: lower jaw in medial aspect. Abbreviations: ar: articular; dc: coronoid process; de: dentary; me: Meckel s cartilage; re: retroarticular. Scale bar: 1 mm. 31 aqua vol. 11 no

34 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: metapterygoid; 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. Neurosensory network: Sensory canals developed on nasal, frontal, pterotic, posttemporal, preopercle, infraorbital, dentary and articular bones, lateral line absent. Infraorbital series comprising four elements. Lacrimal (infraorbital 1) horizontal axis elongate, located superficial to premaxilla (when mouth closed), bearing facets for articulation with lateral ethmoid, posterior border with recess contacting lateral ethmoid lacrimal condyle (Fig. 3). Lacrimal medial face bearing osseous shelf, coursing ventral to nasal sac. Second infraorbital planar, irregular. Third infraorbital cylindrical. Dermosphenotic at posterolateral margin of orbit, contacting sphenotic postorbital process. Supraorbital canal coursing antero-posteriorly from nasal medial margin, along frontal sulcus, curving laterally to confluence with temporal and infraorbital canals. Temporal and preopercular canals not conjoined. Mandibular sensory canal coursing along ventral border of articular and dentary. Canal openings on dentary ventral surface. Hyobranchial apparatus: Comprising median unpaired glossohyal, basihyal, three basibranchials and urohyal; and bilaterally paired dorsal and ventral hypohyals, anterior and posterior ceratohyals, interhyal, six branchiostegal rays, three hypobranchials, five ceratobranchials, four epibranchials and three pharyngobranchials (Fig.10). Basihyal rhomboidal, ossified posteriorly, strongly attached to hypohyals. Basihyal cartilaginous anterior portion supporting small tooth plate. Urohyal with bilateral posterodorsal processes and median posteroventral process. Three median basibranchials coursing antero-posteriorly from basihyal to irregular cartilaginous nodule. Posteriormost two basibranchials fused dorsally to respective tooth plates. Hyoid bar comprising dorsal and ventral hypohyals, and anterior and posterior ceratohyals. Hyoid bar joined to contralateral structure by strong ligament from medial face of hypohyals, coursing ventral to basihyal. Anterior and posterior ceratohyals joined about dentate suture. Anterior ceratohyal ventral surface giving origin to two anteriormost small bran- aqua vol. 11 no

35 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. Abbreviations: an: anterior ceratohyal; at: urohyal anterodorsal process; b1-3: basibranchials 1-3; bh: basihyal; br: branchiostegal ray; bt: free branchiostegal toothplate; c1, c5: ceratobranchials 1 and 5; ch: ceratohyal; cn: cartilage nodule; cr: cranial 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: urohyal posteroventral process; un: epibranchial uncinate process; vh: ventral hypohyal. Scale bar: 1 mm. 33 aqua vol. 11 no

36 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

37 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 posteromedial 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

38 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. Abbreviation: ps: postmaxillary process. Scale bar: 1 mm. Fig. 16. Iso natalensis, MU I-227, 40.1 mm SL. Premaxilla. Abbreviation: ps: postmaxillary process. Scale bar: 1 mm. aqua vol. 11 no

39 Basim Saeed, Walter Ivantsoff and Aarn Fig. 17. Iso natalensis, MU I-227, 45.1 mm SL. Scanning electron micrograph of premaxilla. Fig. 18. Iso hawaiiensis, BPBM 10012, 28 mm SL. Scanning electron micrograph of neurocranium in ventral 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. Abbreviations: ap: pterosphenoid anterior process; fr: frontal; mp: pterosphenoid medial process; ps: pterosphenoid; sp: sphenotic. chiostegal rays, then two large rays rising from anterior 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, ceratobranchial, epibranchial and pharyngobranchial, fourth arch of ceratobranchial and epibranchial, fifth arch comprising single ceratobranchial. Gill rakers of Fig. 20. Notocheirus hubbsi, MU I-307, 32.3 mm SL. Cranium in dorsal aspect. Abbreviations: ec: ethmoid cartilage; ep: epiotic; fr: frontal; le: lateral ethmoid; pi: parietal; po: postorbital process; so: supraoccipital; sp: sphenotic; tc: temporal canal; vo: vomer. 37 aqua vol. 11 no

40 Descriptive anatomy of Iso rhothophilus (Ogilby), with a phylogenetic analysis of Iso and a redefinition of Isonidae (Atheriniformes) first gill arch elongate. Ceratobranchials elongate, arcuate, ventral surface bearing process for attachment of pharyngeal musculature. Fifth ceratobranchial posterolateral angle produced as site of attachment for posterior levator muscle. Hypobranchials fused dorsally to respective tooth plates. Basibranchial tooth plate dorsal to, but not fused with, third hypobranchial. Tooth plates adjacent to, but not fused with, first four ceratobranchials and all epibranchials. Fifth ceratobranchial fused to large lower pharyngeal toothplate. Axial skeleton: Total vertebrae 38-45, precaudal 14-18, caudal First vertebra with reduced neural spine (Fig. 11), articulating with cranium. Neural arch rising from anterodorsal portion of respective vertebral centrum. Neural arch of vertebra 2-6/7 developed as expanded neural plate. Precaudal vertebrae with parapophyses rising from anteroventral portion of centrum. Anterior parapophyses small, directed laterally, anteroposterior series becoming larger, directed posteriorly to form haemal arches of caudal vertebrae. First haemal arch emanating from first caudal vertebra, anterior parapophysis directed ventromedially to fuse with contralateral structure, produced as haemal spine. Total pleural ribs 12-14, first rib contacting third vertebra. Epural articulating with respective vertebral body posterodorsal to base of parapophysis. First epipleural contacting dorsolateral aspect of parapophysis of third vertebra, coursing posterodorsally to ribs. Intervertebral articulations mediated by zygapophyses. Dorsal zygapophyses prominent on anterior vertebrae, ventral zygapophyses present on posterior precaudal and caudal vertebrae. Penultimate vertebra bearing very large haemal spine. Terminal half-centrum supporting hypurals and parhypural. Parhypural fused with hypurals 1 and 2, separated by horizontal gap from fused hypurals 3 and 4. Uroneural coursing posterodorsally from termi- Fig. 21. Iso flosmaris, SOSC VGS-68-30, 52 mm SL. Scanning electron micrograph of posterior neurocranium in dorsal aspect. Abbreviations: ep: epiotic; fr: frontal; pc: epiotic crest; pi: parietal; so: supraoccipital. Fig. 22. Iso hawaiiensis, BPBM 10012, 28 mm SL. Scanning electron micrograph of portion of neurocranium. Abbreviations: eo: exoccipital; ic: intercalar; 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

41 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 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 I. flosmaris I. nesiotes I. rhothophilus 2 3 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

42 Descriptive anatomy of Iso rhothophilus (Ogilby), with a phylogenetic analysis of Iso and a redefinition of Isonidae (Atheriniformes) nal half-centrum, epural anterior to uroneural. Fins and girdles: Each spine of first dorsal supported by individual basal radial comprising fused proximal and medial elements. No radials between dorsal fins. Each element of second dorsal and supported by basal radial and non-ossified distal radial, basal radials also supporting preceding element. Pectoral fin with four proximal radials supporting fin rays, dorsal radial fused to scapula (Fig. 6A). Scapular foramen between scapula and cleithrum. Coracoid contacting cleithrum and scapula dorsally, ventrally Fig. 25. Notocheirus hubbsi, CAS 5526 (paratype). Pectoral girdle. Abbreviations: cl: cleithrum; co: coracoid; pc: postcleithrum; po: posttemporal; ra: proximal radial; sc: scapular; sf: scapular foramen. Scale bar: 1mm. forming medial shelf and additional contact with cleithrum in most specimens. Supracleithrum absent. Posttemporal trabecular, arcuate, superficial to epiotic dorsal crest. Dorsal postcleithrum absent, ventral postcleithrum contacting coracoid. Pelvic fin without distal radials, fin elements supported directly (Fig. 12). Adductor profundus pelvicus muscle on dorsomedial aspect of pelvic, abductor superficialis on lateral aspect. Pelvic posteromedial process elongate, extending dorsally. Pelvic posteroventral process fused to contralateral structure. 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 discontinuous; 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 = symphysis 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 external 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). 8. Basihyal teeth. There are no teeth on the basihyal of Notocheirus and I. flosmaris. Other taxa have teeth on the basihyal (Fig. 10). aqua vol. 11 no

43 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 (0 = basihyal teeth absent; 1 = teeth present). 9. Urohyal posterodorsal process. In all taxa the urohyal posterodorsal process is directed posterodorsally (Figs. 10B, 23) except in I. nesiotes, in which the process is directed dorsally (Saeed et al., 1993; Figs. 2Ca, 2Cb). (0 = urohyal posterior process directed posterodorsally; 1 = process directed dorsally). 10. Anterior vertebral dorsal postzygapophyses. All taxa have well-defined processes (Fig. 11) except I. hawaiiensis (Fig. 24) in which the processes are not discernable. (0 = anterior vertebral dorsal postzygapophyses well defined; 1 = processes not defined). 11. Coracoid. The coracoid is relatively small in Notocheirus (Fig. 25) and I. nesiotes (Saeed et al., 1993; Figs. 2Da, 2Db), and large in other taxa. (0 = coracoid small; 1 = coracoid large). Eleven characters thus identified were entered into a data matrix (Table I). A second matrix was then developed to show the number of shared plesiomorphies between pairs of taxa (Table II). This was used to generate the cladogram (Fig. 26). Discussion The hierarchy of relationships in Iso was I. flosmaris (I. nesiotes (I. rhothophilus (I. hawaiiensis, I. natalensis))). This scheme postulates 5 reversals, while providing deep analysis of the genus. Said (1983) examined five species (and a number of putative hybrid forms), and used cluster and discriminant function analyses of osteological and/or morphological and/or meristic data, to examine relationships within Iso. Although no two hierarchies were identical, the majority of schemes supported (I. flosmaris ((I. rhothophilus, I. natalensis), (I. hawaiiensis, I. nesiotes))). The present scheme, based on a phylogenetic approach, supports the position of I. flosmaris as a sister group to the other four species, but is otherwise dissimilar. Isonidae was erected by Rosen (1964) with eleven osteological characters. Said (1983) described 33 morphological and/or anatomical characters differing between Iso and Notocheirus. Subsequently Saeed et al. (1994) erected Atherinopsoidea, comprising 41 aqua vol. 11 no

44 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. (Atherinopsidae, Isonidae, Notocheiridae), with five osteological characters, distinguishing this group from Atherinoidea. Notocheiridae was diagnosed with seven characters, while Isonidae was diagnosed with six characters. Dyer and Chernoff (1996) considered Iso and Notocheirus to be more derived than Atherinopsidae, and a sister group to the so-called infraorder Atherines (comprising four families Melanotaeniidae, Atherionidae, Phallostethidae and Atherinidae). On the basis of examination of I. natalensis, I. rhothophilus and N. hubbsi, and consideration of previous studies, (Iso, Notocheirus) Notocheiridae was diagnosed with three autapomorphies: supracleithrum absent; pelvic dorsolateral process elongate, attaching to pleural rib by elongate ligament; and body shape unique. Iso was diagnosed with six autapomorphies and four additional characters, Notocheirus was diagnosed with seven autapomorphies and six additional characters. Thus, Dyer and Chernoff (1996) identified 23 anatomical differences between Iso and Notocheirus. Table III lists twenty autapomorphies of either Iso or Notocheirus the present authors have been able to verify, following Rosen (1964), Said (1983), Saeed et al. (1994) and Dyer and Chernoff (1996). Some of the character states in Notocheirus (including lack of first dorsal, mesethmoid, first epibranchial and epural) could be interpreted as paedomorphism (eg. Weitzmann and Vari, 1988), and hence advanced rather than plesiomorphic traits. In any event, the distinction between Notocheiridae and Isonidae is supported by 20 characters. Notocheirus is known only from coastal waters of South America, a region in which Iso has never been collected. It is not known whether the separation between Iso and Notocheirus commenced before the land masses of present day Africa and the Americas were conjoined. It is to be hoped that comparisons of molecular sequences will advance the present knowledge of isonid systematics. Acknowledgements We wish to thank Ms Betty Thorn and Mr. Ron Oldfield (Macquarie University) for preparation of some illustrations and micrographs, and staff of institutions who kindly facilitated loans of specimens. References Clark, H. W. (1937). New fishes from Templeton Crocker Expedition of Copeia, 2: Dyer, B. S. & B. Chernoff. (1996). Phylogenetic relationships among the atheriniform fishes (Teleostei, Atherinomorpha). Zoological Journal of the Linnaean Society, London, 117: Golvan, Y. J Catalogue systématique des noms de genres de poissons actuels, de la 10 e édition Systema naturae de Charles Linné jusqu à la fin de l année Masson et Cie, Paris 227pp. Herre, A. W Notes on the fishes in the Zooological Museum of Stanford University. XVII. New fishes from Johore and India. Proceeding of the Biological Society of Washington, 57: Jordan, D. S. & C. L. Hubbs Studies in Ichthyology. A monographic review of the family Atherinidae or silversides. Leland Stanford Junior University Publications, University Series. 87 pp. Jordan, D. S. & E. C. Starks. (1901). A review of the atherine fishes of Japan. Proceedings of the United aqua vol. 11 no

45 States National Museum. 24: McCulloch, A. R A check-list of the fishes recorded from Australia. Memoirs of the Australian Museum, 5 (1): Ogilby, J. D On two genera and species of fishes from Australia. Proceedings of the Linnean Society of N.S.W. 10: Regan, C. T fishes from Durban collected by Messrs. H. W. Bell Marley and Romer Robinson. Annals of Durban Museum, 2 (4): 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: Saeed, B., Ivantsoff, W. & G. R. Allen Taxonomic revision of the family Pseudomugilidae (Order Atheriniformes). Australian Journal of Marine and Freshwater Research, 40: Saeed, B., Ivantsoff, W. & L. E. L. M. Crowley A new species of the surf-inhabiting Atheriniform Iso (Pisces: Isonidae). Records of the Western Australian Museum. 16: Saeed, B., Ivantsoff, W. & L. E. L. M. Crowley Systematic relationships of atheriniform families within division 1 of the series Atherinomorpha (Acanthopterygii) with relevant historical perspectives. Journal of Ichthyology, 34 (9): Said, B. M Revision of the fish genus Iso. Unpublished MSc thesis, Macquarie University, Australia, 177 pp. Said, B Revision of the genus Pseudomugil with phylogenetic systematics of the order Atheriniformes. Unpublished. Ph.D. thesis, Macquarie University, Australia, 280 pp. Schultz, L. P 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: Waite, E. R. 1904b. Synopsis of the fishes of New South Wales. Memoirs of the New South Wales Naturalist Club, 2:1-59. Weitzman, S. H. & R. P. Vari Miniaturization in South American freshwater fishes; an overview and discussion. Proceedings of the Biological Society of Washington, 101: Winterbottom, R A descriptive synonymy of the striated muscles of the Teleostei. Proceedings of the Academy of Natural Sciences of Philadelphia, 125: Basim Saeed, Walter Ivantsoff and Aarn 43 aqua vol. 11 no

46 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 Allen, Gerald R. and John E. Randall: A new species of damselfish (Pomacentrus: Pomacentridae) from Fiji. aqua 10 (3): , October 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 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): , October 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 Møller, Peter Rask, Werner Schwarzhans and Jørgen G. Nielsen: Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia. aqua 10 (4): , November 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 Randall, John E.: Chlorurus perspicillatus x C. sordidus, a hybrid parrotfish from the Hawaiian Islands. aqua 10 (1): 39-43, June 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 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 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 Winterbottom, Richard: Two new species of the Trimma tevegae species group from the Western Pacific (Percomorpha: Gobiidae). aqua 10 (1): 29-38, June Winterbottom, Richard: Feia dabra, a new species of gobiid fish (Percomorpha: Gobiidae) from Palau. aqua 10 (2): 45-50, July Winterbottom, Richard: On the Status of Trimma tevegae and Trimma caudomaculata (Percomorpha: Gobiidae). aqua 10 (2):51-56, July Zuanon, Jansen and Ivan Sazima: The ogre catfish: prey scooping by the auchenipterid Asterophysus batrachus. aqua 10 (1): 15-22, June 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 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 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 Feia dabra n. sp. a new species of gobiid fish (Percomorpha: Gobiidae) from Palau. aqua 10 (2): 45-50, July 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 Ogilbia boehlkei n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia. aqua 10 (4): , November Ogilbia boydwalkeri n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia. aqua 10 (4): , November Ogilbia cocoensis n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia. aqua 10 (4): , November Ogilbia davidsmithi n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia. aqua 10 (4): , November Ogilbia jeffwilliamsi n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia. aqua 10 (4): , November Ogilbia jewettae n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia. aqua 10 (4): , November Ogilbia mccoskeri n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia. aqua 10 (4): , November Ogilbia nigromarginata n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia. aqua 10 (4): , November Ogilbia nudiceps n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia. aqua 10 (4): , November Ogilbia robertsoni n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia. aqua 10 (4): , November Ogilbia sabaji n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia. aqua 10 (4): , November Ogilbia sedorae n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia. aqua 10 (4): , November Ogilbia suarezae n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia. aqua 10 (4): , November 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 Ogilbia tyleri n. sp. Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia. aqua 10 (4): , November Pomacentrus microspilus n. sp. A new species of damselfish (Pomacentrus: Pomacentridae) from Fiji. aqua 10 (3): , October 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 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 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 A synopsis of coral and coral-rock associated gobies (Pisces: Gobiidae) from the Gulf of Aqaba, northern Red Sea. aqua 10 (3): , October Book review: Freshwater Fishes of north-eastern Australia. aqua 10 (4): 208, November Book review: Reef and Shore Fishes of the South Pacific New Caledonia to Tahiti and the Pitcairn Islands. aqua 10 (3):102, October Chlorurus perspicillatus x C. sordidus, a hybrid parrotfish from the Hawaiian Islands. aqua 10 (1): 39-43, June On the Status of Trimma tevegae and Trimma caudomaculata (Percomorpha: Gobiidae). aqua 10 (2):51-56, July Review of the American Dinematichthyini (Teleostei: Bythitidae). Part II. Ogilbia. aqua 10 (4): , November The ogre catfish: prey scooping by the auchenipterid Asterophysus batrachus. aqua 10 (1): 15-22, June The sunfish Mola mola as an attachment surface for the Lepadid Cirriped Lepas anatifera a previously unreported association. aqua 10 (1): 1-4, June aqua vol. 11 no

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48 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 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 Gerald R. Allen, Forrest Young and Patrick L. Colin: Centropyge abei, a new species of deep-dwelling angelfish (Pomacanthidae) from Sulawesi, Indonesia John E. Randall and John L. Earle: Amblyeleotris neumanni, a new species of shrimp goby from New Britain Basim Saeed, Walter Ivantsoff and Aarn: Descriptive anatomy of Iso rhothophilus (Ogilby), with a phylogenetic analysis of Iso and a redefinition of Isonidae (Atheriniformes) Index aqua 10(1-4) Papers appearing in this journal are indexed in: Zoological Record; Biolis - Biologische Literatur Information Senckenberg; 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 culture in aqua Vol. 11 (2). Photo by E. K. Balon.

A new killifish of the genus Melanorivulus from the upper Paraná river basin, Brazil (Cyprinodontiformes: Rivulidae)

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