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aquaJournal of Ichthyology and Aquatic

BiologyVol. 7 (1), May 2003

AquapressISSN

0945-9871

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Managing Editor:

Heiko BleherVia G. Falcone 11, 27010 Miradolo Terme (PV), ItalyTel.: +39 0382 754707/08 - Fax: +39 0382 754129e-mail: [email protected]

Scientific Editor:

Dr. Walter IvantsoffSenior Research Fellow,Department of Biological Sciences,Macquarie University, N.S.W. 2109, AustraliaTel. +61 2 9850 8167 - Fax +61 2 9869 8886 e-mail: [email protected]

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 94720-3140,U.S.A.

Henri J. Dumont, Rijksuniversiteit Gent, Laboratoriumvoor Ecologie der Dieren, Zoogeografie en Natuur-behoud, K. L. Ledeganckstraat, 9000 Gent, Belgium

Jacques Géry, Chemin du Plantier, 24200 Sarlat,France

Frank Kirschbaum, Institut für Gewässerökologie undBinnenfischerei, Abt. 4 Forschungsverbund Berlin e. V.Müggelseedamm 310, 12587 Berlin, Germany

Friedhelm Krupp, Forschungsinstitut Senckenberg,Senckenberganlage 25, 60325 Frankfurt am Main,Germany

Christian Lévêque, CNRS - Programme EnvironnementVie et Sociétès, 1 Place Aristide Briand, 92195 ParisCédex, France

Volker Mahnert, Muséum d’Histoire Naturelle, Route deMalagnou 1, 1211 Genève 6, Switzerland

Robert R. Miller, University of Michigan, Museum ofZoology, Ann Arbor, Michigan 48109, U.S.A.

Paolo Parenti, Department of Enviromental Sciences,University of Milan-Bicocca, Piazza della Scienza 1, I-20126 Milan, Italy

John E. Randall, Bishop Museum, 1525 Bernice Street,P.O. Box 19000-A, Honolulu, Hawaii, U.S.A.

Wolfgang Schneider, Hessisches Landesmuseum,Darmstadt, Friedensplatz 1, 64283 Darmstadt, Germany

Lothar Seegers, Grenzstraße 47b, 46535 Dinslaken,Germany

Wolfgang Villwock, Universität Hamburg, ZoologischesInstitut und Zoologisches Museum, Martin-Luther-King-Platz 3, 20146 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 originalscientific articles in the fields of systematics, taxonomy ,biogeography, ethology, ecology, and general biology offishes, amphibians, aquatic invertebrates, and plants.Papers on freshwater , brackish, and marine organismswill be considered. aqua is fully refereed and aims atpublishing manuscripts within 2-4 months of acceptance.With the publication of aqua we are pursuing a new con-cept: In view of the importance of colour patterns inspecies identification and animal ethology , authors areencouraged to submit colour illustrations as well asdescriptions of coloration. It is our aim to provide the international scientific community with an ef ficientlypublished series meeting high scientific and technicalstandards.

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The editors welcome the submission of original manu-scripts which should be sent directly to the scientific editor. Full length research papers and short notes will beconsidered for publication. There are no page chargesand colour illustrations will be published free of charge.Authors will receive 50 free reprints of each paper .

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ISSN 0945-9871Publisher: Aquapress, Redazione aqua, I-27010 Miradolo Terme (Pavia), ItalyPrinter: S.A.T.E. s.r.l., (Bergamo) ItalyTypesetting: Rossella Bulla© 2003 aqua, Journal of Ichthyology and Aquatic Biology

aqua - Journal of Ichthyology and Aquatic Biology

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KeywordsTaxonomy, labrid fishes, Thalassoma, new species,

south-west Pacific

Abstract Thalassoma nigrofasciatum is described as a new

species of Labridae from the Great Barrier Reef, V anu-atu, Lord Howe Island, Norfolk Island, ChesterfieldBank, New Caledonia, Loyalty Islands, Fiji, and Tonga.It is typically found on exposed outer-reef areas fromtide pools to depths of at least 15 m. Previously identi-fied as T. jansenii (Bleeker), it dif fers in colour and inhaving a deeper caudal peduncle and longer paired fins.

ZusammenfassungThalassoma nigrofasciatum wird als neue Art der

Familia Labridae vom Großen Barrierenrif f, von Vanu-atu, von der Lord-Howe-Inselgruppe und der Norfolk-Insel, Cesterfield Bank, Neu-Kaledonien, der Loyalty-Inselgruppe, aus Fidschi und Tonga beschrieben. Typ-ischerweise wird diese Art in of fenen, ungeschütztenRiffgebieten, von Tidentümpeln bis hinunter zu einerTiefe von mindestens 15 Metern gefunden. Vormals alsT. jansenii (Bleeker) identifiziert, unterscheidet sie sichanhand ihrer Färbung, sowie durch einen breiterenSchwanzstiel und durch längere, paarige Flossen.

RésuméThalassoma nigrofasciatum est décrit en tant qu'e-

spèce nouvelle de labridé, originaire de la Grande Bar-rière de Corail, V anuatu, l'île Lord Howe, l'île Norfolk,Chesterfield Bank, la Nouvelle-Calédonie, les îles Loy-auté, les îles Fidji et Tonga. Le biotope typique consisteen zones externes et exposées de récifs, depuis lesflaques de marées jusqu'à des profondeurs d'au moins15 m. D'abord identifiée comme T. jansenii (Bleeker),l'espèce se distingue par la couleur , par un pédonculecaudal plus large et par des nageoires paires pluslongues.

SommarioThalassoma nigrofasciatum viene descritta come

nuova specie di Labridae sulla base di esemplari rac-colti lungo numerose località del Pacifico sudocciden-tale, tra cui la Grande Barriera Australiana, Vanuatu, le

isole Lord Howe e Norfolk, Chesterfield Bank, NuovaCaledonia, Isole della Lealtà, Fiji e Tonga. Si localizzatipicamente sulla parte esterna della barriera in acquesuperficiali fino a 15 m di profondità. Precedentementeidentificato come T. jansenii (Bleeker), ne differisce perla colorazione, il peduncolo caudale più alto e le pinnedoppie più allungate.

IntroductionHeiser (1981, unpublished PhD thesis) recognized

21 species of labrid fishes in the genus ThalassomaSwainson, of which 14 are Indo-Pacific species (oneas Thalassoma sp.). He overlooked T. septemfascia-tum Scott from Western Australia and failed to recog-nize T. tri lobatum (Lacépède). Randall and Edwards(1994) reviewed the complex of six species thatincludes trilobatum and described T. heiseri from thePitcairn Islands and Tuamotu Archipelago (the Tha-lassoma sp. of Heiser, 1981). Randall and Mee (1994)described T. loxum from O man. R andall ( 1995)reviewed the three eastern Pacific species of thegenus and showed that records of T. lutescens (Layand Bennett) from the Galápagos are misidentifica-tions of T. grammaticum Gilbert. Allen (1995) addeda fourth eastern Pacific species when he described T.robertsoni from Clipperton Island. Fricke (1999)described T. mascarenum from the MascareneIslands, but Randall and Smith (2001) placed it in thesynonymy of T. genivittatum (Valenciennes).

Bleeker (1856) described Julis jansenii (now classi-fied in Thalassoma) as a new species of labrid fishfrom eight specimens, 90-165 mm SL, collected atManado, Sulawesi. In the first volume of his AtlasIchthyologique (1862), he questioned whether Julisblochii Valenciennes in Cuvier and V alenciennes(1839), reported from the East Indies, might be asenior synonym of Julis jansenii . Valenciennes haddetermined that the species illustrated by Bloch(1791) as Labrus lunaris Linnaeus could not be thatspecies and provided the new name Julis blochii .Heiser (1981) concluded that neither Bloch’ s plate orhis description of Labrus lunare (Valenciennes onlydescribed the colour of blochii from Bloch’s plate) pro-vided enough information for positive identification asany Thalassoma we know today. Paepke (1999) listed

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aqua, Journal of Ichthyology and Aquatic Biology

Thalassoma nigrofasciatum, a new species of labrid fish from the south-west Pacific

John E. Randall

Bishop Museum, 1525 Bernice St., Honolulu,Hawai’i 96817-2704, USA

Accepted: 29.03.2003

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all.

two syntypes of Julis blochii in the fish collection of theMuseum für Naturkunde der Humboldt Universität inBerlin as ZMB 2579. The author asked Peter Bartschof this Museum for information on the syntypes. Ran-dall and Bartsch (MS) selected the larger of the twospecimens (the one illustrated by Bloch) as the lecto-type of Julis blochii. They determined that Julis blochiiis a junior synonym of Thalassoma pavo (Linnaeus)from the eastern Atlantic. Bloch’s locality of the EastIndies was an error.

The author and others have long been aware ofcolour variation in Thalassoma jansenii and specu-lated on the possibility of there being two species. Inthe Indo-Malayan region north to Japan, east to theMariana and Caroline Islands, and west to the Mal-dive Islands, the body of the initial phase has five orsix broad black bars separated by narrow white topale yellow bars, whereas in the South Pacific fromthe Great Barrier Reef to Fiji and Tonga there are justthree broad black bars, the anterior continuing ontothe postorbital head. The terminal-male phase of thetwo is more similar, but the middle black bar reachesfarther ventrally on the south-west Pacific form, andthe posterior edge of this bar is vertical or nearly so.An early attempt to find some meristic or morphologi-cal dif ferences to correlate with the colour patternswas not unsuccessful, and in a more determinedrecent ef fort to find morphological dif ferences, the

length of the pectoral fins, and to a lesser extent thepelvic fins, was found to be shorter in jansenii, and thedepth of the caudal peduncle not as deep as injansenii.

Logistic difficulties did not permit an electrophoreticstudy on frozen specimens as planned in 1993.

The South Pacific form is described here as new, thusraising the total number of species for Thalassoma to27, making it the third most speciose genus of the family Labridae (after Halichoeres and Cirrhilabrus).

Materials and MethodsType specimens of the new species have been

deposited in the Australian Museum, Sydney (AMS);Bernice P. Bishop Museum, Honolulu (BPBM), Cali-fornia Academy of Sciences, San Francisco (CAS;SU); and the National Museum of Natural History ,Washington, D.C. (USNM).

Lengths of specimens are given as standard length(SL), measured from the most anterior end of theupper lip to the base of the caudal fin (posterior end ofhypural plate); head length is measured from the sameanterior point to the posterior end of the opercular flap;body depth is taken vertically from in front of the baseof the pelvic fins; body width is the maximum width justposterior to the gill opening; orbit diameter is the great-est fleshy diameter , and interorbital width the leastfleshy width; upper jaw length is taken from the front of

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Table I. Proportional measurements of type specimens of Thalassoma nigrofasciatum expressed as percentages of thestandard length.

Holotype ParatypesBPBM BPBM BPBM CAS BPBM BPBM BPBM BPBM14768 31161 33719 40164 31742 11351 14769 14769

Standard length (mm) 131.8 58.2 62.2 74.4 81.0 94.0 116.1 132.0Body depth 30.1 30.5 28.8 29.6 29.7 29.3 28.0 29.3Body width 14.4 14.2 16.9 13.4 16.4 12.7 12.6 13.2Head length 30.6 32.1 30.6 32.2 29.3 32.4 31.0 29.4Snout length 11.8 10.2 10.6 11.1 10.5 11.6 11.1 11.9Orbit diameter 5.0 7.4 7.2 6.8 6.9 6.4 6.0 5.3Interorbital width 9.1 8.6 8.1 8.5 8.6 8.8 10.5 8.8Caudal peduncle depth 15.2 15.8 16.1 15.5 15.9 15.0 15.3 15.4Caudal peduncle length 12.8 12.7 12.2 11.9 12.5 12.3 12.6 12.1Upper jaw length 9.1 9.2 8.9 9.4 9.0 9.7 9.8 9.3Predorsal length 36.3 34.8 35.0 36.4 35.6 35.2 38.0 35.6Preanal length 60.0 59.7 58.3 59.2 57.9 60.0 60.1 60.5Prepelvic length 35.7 34.9 34.4 35.4 34.8 36.5 36.1 35.7Dorsal fin base 56.7 58.6 59.2 58.3 59.5 57.6 56.3 57.2First dorsal spine 5.1 6.4 6.4 6.6 5.0 6.3 5.3 4.8Longest dorsal spine 11.6 11.9 12.8 12.5 12.9 11.7 11.5 11.0Longest dorsal ray 13.5 15.0 14.4 14.9 14.8 12.8 12.5 12.3Anal fin base 28.4 30.1 30.9 28.8 31.8 28.9 29.2 27.3Second anal spine 7.1 8.3 8.1 8.2 7.6 7.5 7.1 7.2Third anal spine 10.8 12.5 11.4 11.5 12.4 12.4 11.1 11.1Longest anal ray 14.0 14.6 14.4 14.8 14.0 13.9 14.7 13.7Caudal fin length 36.4 23.2 22.5 23.7 25.1 26.6 27.4 33.9Caudal concavity 19.2 0 1.7 2.8 5.0 7.4 8.3 11.0Pectoral fin length 24.3 23.8 23.9 24.2 24.9 23.4 25.0 24.2Pelvic spine length 9.3 9.5 9.2 8.9 9.4 8.6 8.8 9.0Pelvic fin length 15.4 15.7 15.2 15.2 15.9 15.1 14.5 14.8

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the upper lip to the posterior end of the maxilla; caudalpeduncle depth is the least depth, and caudal-pedun-cle length the horizontal distance between verticals atthe rear base of the anal fin and the caudal-fin base;lengths of fin spines and rays of the median fins aremeasured from their extreme bases; pectoral finlength is the length of the longest ray; pelvic fin lengthis measured from the base of the pelvic spine to the tipof the longest soft ray. Lateral line scale counts do notinclude the last pored scale on the caudal fin base;pectoral ray counts contain the upper rudimentary ray;gill raker counts include all rudiments.

Data in parentheses in the description of the newspecies refer to paratypes (when dif ferent from theholotype). Table I presents 25 measurements of typespecimens as percentages of the standard length.Ratios of proportional measurements in the text aregiven in terms of the standard length, rounded to thenearest 0.05. Preserved specimens of Thalassomavary greatly in the head length because of the dif fer-ent degree of extrusion of the jaws, so short mea-surements such as orbit diameter and snout lengththat are usually expressed in terms of the head lengthare related here to SL.

Comparative material examined Thalassoma jansenii: CAROLINE ISLANDS: Ifaluk

Atoll, CAS 21 15695, 98 mm. P ALAU: Angaur, CAS73080, 3: 63-82 mm; Auluptagel Island, CAS 73078, 2:119-121 mm; Ngaremdiu Reef, CAS 73079, 63.5 mm;Kayangel Atoll, CAS 73944, 59.5 mm. P APUA NEWGUINEA: Madang Province, W ongat Island, CAS65868, 73 mm. INDONESIA: Sangihe Islands, CAS30038, 3: 76-82 mm; SU 29854, 85 mm. PHILIP-PINES: Jolo, SU 26000, 132 mm. Luzon, SU 26001,103 mm. RYUKYU ISLANDS: Okinawa, SU 20949, 2:

109-144 m m. ANDAMAN S EA: Thailand, S imilanIslands, BPBM 22789, 3: 52.8-74 mm. MALDIVEISLANDS: North Malé Atoll, BPBM 33086, 78.5 mm.

Thalassoma nigrofasciatum n. sp. (Figs. 1-3, Table I)

Thalassoma janseni (non Bleeker) Randall, Allen andSteene, 1990: 33, middle fig. (Great Barrier Reef).

Thalassoma janseni (non Bleeker) Francis, 1991:215, fig. 33 (Norfolk Island).

Thalassoma jansenii (non Bleeker) Eichler andMyers, 1997: 341, bottom fig. (no locality).

Thalassoma jansenii (non Bleeker) Laboute andGrandperrin, 2000: 369, lower 2 figs. (New Caledo-nia).

Holotype: BPBM 14768, male, 131.8 mm, Lord HoweIsland, Phillips Point, 7.5 m, spear, W. Deas, 6 Febru-ary 1973.Paratypes: BPBM 1 1351, 94.0 mm, Fiji, V iti Levu,Rat-tail Pass (first pass east of Suva Harbor), 2.5-6 m,rotenone, J. E. Randall, R. D. Randall, M. Gawel, andO. McCausland, 7 August 1971; BPBM 14469, 2:116.1-132.0 mm, Queensland, Great Barrier Reef,Capricorn Group, One Tree Island, west side, surgechannel, 1-2 m, spear , J. E. Randall, 16 January1973; CAS 40164, 74.4 cm, Great Barrier Reef, LizardIsland, of f C asuarina Beach, Australian Museumparty, 10 November 1975; BPBM 31 161, 58.2 mm,Fiji, Malolo Island (17°47’S,177° 13’E), large islet onsouth-west corner, tide pools, 0-0.3 m, rotenone, V. G.Springer et al., 24 May 1982; BPBM 321742, 81.0mm, Coral Sea, Australia, Osprey Reef, west side(13°53’S, 146°32’E), Pelagic Gully, outside reef north

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Fig. 1. Holotype of Thalassoma nigrofasciatum, BPBM 14768, male, 131.8 mm, Lord Howe Island. Photo by the author.

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of entrance to lagoon, 6 m, spear, J. E. Randall, 26January 1987; BPBM 33719, 2: 44.8-62.2 mm, CoralSea, Chesterfield Bank, Ilot du Mouillage, tide pools,0-0.2 m, rotenone, M. L. Kulbicki, J. E. Randall, P . J.Doherty, and C. Goiran, 28 August 1988; USNM323949, 2: 53.2-64.9 mm, Loyalty Islands, OuvéaAtoll, Bagaat Islet (20°37’18”S, 166° 16’8”W), verticalreef wall and small cave in base of wall with coral rub-ble, 15-22 m, rotenone, J. T. Williams, J.-L. Menou,and P. Tirard, 16 November 1991; AMS I.33746-024,75 mm, Great Barrier Reef, Boot Reef (10°2.72”S,144°41.88’E), 6.5 m, rotenone, J. Leis, M.McGrouther, T. Trnski, S. Reader , H. Larson, and M.Westneat, 27 January 1993; USNM 338735, 4: 70.2-131.0 mm, Tonga, Vava’u, Nuku Island, north shore,dead reef top and in channels, (18°42’54”S,174°2’36”W), 1-3 m, rotenone, J. T. Williams, A.Palaki, M. A. McCormick, C. C. Baldwin, E. A. Powers,and D. G. Smith, 19 November 1993; USNM 362177,72.3 mm, V anuatu, Efate Island, south-east side,Enam Bay, chapparal rock, coral, sand in channels(17°48’16”S, 168°30’9”E), 0-2 m, rotenone, J. T.Williams, D. G. Smith, M. McGrouther , R. D. Mooi,and R. Jimmy, 8 May 1997; USNM 363206, 79.4 mm,Vanuatu, Banks Islands, V anua Lava, Port Patteson,off a point to the north of Sola (13°52’25”S,167°32’12”E), surge channels in a high-energy area,1-3 m, rotenone, J. T. Williams, R. D. Mooi, and M.McGrouther, 17 May 1997.

DiagnosisDorsal rays VIII,13; anal rays III,1 1, the first spine,

when detectable, very slender and short; pectoralrays 15-16 (usually 15); lateral line scales 26-27; headnaked, except for a small patch of scales dorsally onopercle; gill rakers 20-23; body depth 5.9-3.6 in SL;head length 3.1-3.4 in SL; snout length 8.4-9.8 in SL;caudal peduncle depth 6.2-6.65 in SL; caudal fin trun-cate in young to strongly lunate with filamentous lobesin adult males; pectoral fins 4.0-4.3 in SL; pelvic fins6.3-6.9 in SL; initial phase white with postorbital head

and anterior body above pectoral-fin base black withan oblique yellow band above dorsal edge of opercle,a black bar over last 4 or 5 dorsal spines continuingventrally across body, its posterior edge vertical, anda broader black bar over last 9 soft rays of dorsal fin,across body, and posterior anal fin; a black spot onfirst 2 membranes of dorsal fin; terminal males withyellow partly replacing the white, a salmon pink patchon chin, and blue ventrally on head and chest. Attainsabout 20 cm.

DescriptionDorsal rays VIII,13; anal rays III,1 1, the first spine,

when detectable, very slender and short; all dorsaland anal rays branched, the last to base; pectoralrays 15 (15-16, usually 15), the uppermost rudimen-tary, the second unbranched; pelvic rays I,5; principalcaudal rays 14, the upper and lower branched; upperprocurrent caudal rays 6, the most posterior seg-mented; lower procurrent caudal rays 5, the most pos-terior segmented; lateral line scales 26 (plus oneslightly larger pored scale on caudal fin base); scalesabove lateral line to origin of dorsal fin 2.5; scalesbelow lateral line to origin of anal fin 8.5; oblique rowsof scales dorsally on nape anterior to origin of dorsalfin 6; circumpeduncular scales 16; branchiostegalrays 6; gill rakers 21 (20-23); pseudobranchial fila-ments 33 (increasing with growth, 14 in 44.8-mmparatype to 35 in 132-mm paratype; dif ficult to counton large specimens); supraneural bones 1; vertebrae9 + 16.

Body depth 3.3 (3.3-3.6) in SL; body width 2.1 (1.8-2.3) in body depth, 5.9-7.95 in SL; head length 3.25(3.1-3.4) in SL; dorsal profile of snout convex, moreso in adults, the snout length 8.5 (8.4-9.8) in SL; orbitdiameter 20 (13.5-18.9) in SL; interorbital space con-vex, the fleshy width 1 1.0 (9.5-12.3) in SL; caudalpeduncle depth 6.6 (6.2-6.65) in SL; caudal peduncledeeper than long, the length 7.8 (7.9-8.4) in SL.

Mouth terminal, the gape slightly oblique; maxillanearly or just reaching below posterior nostril, the

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Fig. 2. Female of Thalassoma nigrofasciatum, about 140 mm TL, Great Barrier Reef. Photo by the author .

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Fig. 3. Female of Thalassoma nigrofasciatum, about 110 mm TL, Fiji. Photo by the author.

Fig. 4. Female of Thalassoma jansenii, about 89 mm TL, Wakatobi Islands, Indonesia. Photo by the author .

Fig. 5. Female of Thalassoma jansenii, about 125 mm TL, Bali, Indonesia. Photo by the author .

Fig. 6. Male of Thalassoma jansenii, about 130 mm TL, Bali, Indonesia. Photo by the author .

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upper jaw length 1 1.0 (10.3-11.2) in SL; lips slightlyfleshy, the lower with a lateral flap extending ventrallyalong side of lower jaw and fitting flush with the surfacedue to a recessed area behind; inner surface of lips pli-cate, the upper with 6 in holotype, the innermost fleshyand papillose; a pair of strong projecting canine teethanteriorly in each jaw, the lower fitting inside the upperwhen jaws closed, followed by 7 (6-9) progressivelyshorter teeth in upper jaw and 1 1 (8-11) in lower j aw;no tooth at corner of mouth; pharyngeal dentition par-tially dissected from largest paratype; each half ofupper pharyngeal bone with 4 anterior-posterior rows ofteeth, none enlarged, the first row of a single bluntlyconical tooth, the second row of 5 bluntly conical teeth,the third row of 7 teeth that are nodular laterally andmolariform medially, and the last row of 7 molariformteeth; median anterior limb of T-shaped lower pharyn-geal with 3 lateral rows of nodular teeth except for amedial bluntly conical tooth at the front; posterior limb

with 3 anterior-posterior rows of nodular to molariformteeth, the last row with a median molar twice the size oflargest remaining teeth, and 5 or 6 teeth to each side.

Anterior nostril a small short membranous tube set ina depression anterior to or slightly above centre ofeye by a distance in holotype of three-fifths orbit diam-eter; posterior nostril a small aperture covered by ananterior flap in front of upper edge of eye, the internar-ial distance about half orbit diameter in holotype.

Free margin of preopercle very short, consisting onlyof a rounded corner and a vertical edge reaching tolevel of rictus; gill membranes broadly attached toisthmus, with a free fold across.

The 44.8-mm paratype with 12 pores from behindcentre of eye to below front of eye with a short branchextending outward from each and bearing smallpores; with growth, the branches increase in lengthand the pores become more numerous, with as manyas ten on one branch; sensory canal of preopercle

Fig. 7. Male of Thalassoma jansenii (after Bleeker, 1862)

Fig. 8. Female of Thalassoma jansenii, about 110 mm TL, Maldive Islands. Photo by the author .


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with about 13 pore-bearing branches that radiate ven-trally or posteriorly.

Lateral line continuous, angling sharply downwardbelow last 3 rays of dorsal fin to straight peduncularpart; lateral line scales generally with 3 sensorytubules, the adults with short side branches fromthese; juveniles with 2 or 3 pores to each scale, adultswith as many as 15; head naked except for a smallpatch of 5 (4-6) small scales on opercle just anteriorto upper end of gill opening; median predorsal scalesjust reaching a vertical in line with posterior margin ofpreopercle; smallest scales of nape and chest lessthan half size of largest scales of side of body; dorsaland anal fins with a low basal scaly sheath; smallscales on base of middle of caudal fin less than halfdistance to posterior margin, but farther out on lobes;no scales on pectoral fins; a midventral row of 3 smallscales at base of pelvic fins.

Origin of dorsal fin above second lateral line scale,the predorsal length 2.75 (2.65-2.9) in SL; first dorsalspine 19.6 (15.2-20.8) in SL; dorsal spines progres-sively longer posteriorly , the eighth 8.6 (7.75-9.1) inSL; first and second dorsal soft rays longest, 7.4(6.65-8.15) in SL (rays relatively shorter with growth);origin of anal fin below base of third dorsal soft ray,the preanal distance 1.65 (1.65-1.75) in SL; first analspine very short and slender , often not detectablewithout dissection or X-ray; second anal spine 14.1(12.1-14.1) in SL; third anal spine 9.25 (8.0-9.0) in SL;first and second anal soft rays longest, 7.15 (6.75-7.3)in SL; caudal fin truncate in juveniles, lunate with pro-longed lobes in adults, the fin length 2.75 (2.95-4.45)in SL, the caudal concavity 5.2 (0-9.1) in SL; third andfourth pectoral rays longest, 4.1 (4.0-4.3) in SL; originof pelvic fins below lower base of pectorals, the pre-pelvic length 2.8 (2.75-2.9) in SL; first pelvic soft raylongest, 6.5 (6.3-6.9) in SL

Colour when live: Colour of holotype (male) whenfresh is shown in Fig. 1. Larger males seen in thesame area could be distinguished by more blue andmore dusky pigment in the white sections of the headand body, more yellow on the vertical pale bar in mid-dle of body, and blackish lobes of the caudal fin.

The colour in life of a probable large female of about140 mm total length from the Great Barrier Reef maybe seen in Fig. 2. The colour in life of a female ofabout 100 mm total length is shown in Fig. 3.

Colour in alcohol: Colour of holotype, an adultmale: body pale yellowish with an oblique dark brownband from nape to beneath basal half of pectoral fin; adark brown bar extending across body from base oflast 5 spines of dorsal fin, narrowing as it passes ven-trally, its posterior edge nearly vertical and ending atanus; a broad dark brown bar covering all of dorsal finposterior to third soft ray, crossing body, and all of analfin posterior to second soft ray , the posterior marginpassing between first and second peduncular lateralline scales; head pale yellowish below level of lower

edge of eye; snout above upper lip and interorbitalbrownish gray; postorbital head dark brown with anoblique pale yellowish band behind opercular mem-brane above base of pectoral fin; dorsal fin anterior tofifth spine dark brown, with a black spot on secondmembrane; caudal fin brownish gray with a broad paleyellowish semicircular area midposteriorly in fin, therays within brown; paired fins pale yellowish, the pec-torals with a small triangular dark brown spot at upperbase that extends more broadly onto axil of fins.

A 44.8-mm juvenile from the Coral Sea is pale yellowish in preservative with two broad dark brownbars covering most of body, the first ending beneathpectoral fin, the two bars separated by a pale bar oforbit diameter in width on midside of body andexpanded dorsally and ventrally; dark bars extendingonly faintly into base of dorsal and anal fins; a blackspot on each of first two membranes of dorsal fin, anda small dark spot at rear base of last ray of dorsal fin;all fins pale.

EtymologyThis species is named nigrofasciatum from the Latin,

in reference to the presence of black bars on the body.

RemarksThalassoma nigrofasciatum is known from the Great

Barrier Reef from Boot Reef at 10° north to the Capri-corn Group in the south, Lord Howe Island, NorfolkIsland, Kermadec Islands (Francis, 1993), New Cale-donia, Loyalty Islands, Chesterfield Bank in the CoralSea, Fiji, and Tonga. It is typically found inshore inouter reef areas; collections were made from tidepools to a depth of 15 m.

As discussed in the introduction, this species hasbeen considered as a colour variant of Thalassomajansenii (Bleeker), described from Sulawesi. Typical T.jansenii is known from the Mariana Islands, CarolineIslands, Palau, Indonesia, Philippines, Taiwan, south-ern Japan, Andaman Sea, Christmas Island (easternIndian Ocean), and W estern Australia. Juveniles andfemales of jansenii have five or six black bars dorsallyon the body , in addition to the one on the postorbitalhead, separated totally or partially by narrow white topale yellow bars (Figs. 4, 5); in males the bars coalesceand extend farther ventrally, such that the pattern mayresemble that of a male nigrofasciatum, but the middleblack bar does not extend as far ventrally , and its pos-terior edge is not vertical (see Figs. 6 and 7, the latterafter Bleeker , 1862). An exception is CAS 65868, afemale 73 mm SL from Madang Province, Papua NewGuinea, in which there is a vertical demarcation in themiddle of the body, but the dark brown anterior part ofthe body is continuous with the dark postorbital head,and the black portion in the middle of the body reachesonly about three-fourths of the body depth.

Thalassoma nigrofasciatum differs from jansenii inhaving a deeper caudal peduncle, its depth in 15 type

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specimens 14.9-16.2% SL (mean 15.4%), comparedto 16 specimens of jansenii the same approximatesize, 14.2-15.1% (mean 14.6%). The pectoral fins ofnigrofasciatum are longer , 23.4-25.7% SL (mean24.5%) compared to 21.6-23.7% (mean 22.6%) forjansenii, and the pelvic fins are longer on the average,14.5-16.9% SL (mean 15.3%), compared to 13.3-15.3% (mean 14.5%) for jansenii. The above-men-tioned specimen from Papua New Guinea has pedun-cular and paired-fin measurements that are modal fornigrofasciatum.

One specimen of Thalassoma jansenii from the Mal-dive Islands (BPBM 33086, 78.5 mm), the only oneexamined, has the least caudal peduncle depth,14.0% SL, and the shortest paired fins, the pectorals21.8% SL, and the pelvics 12.8% SL. It was notincluded in computing the percentages for janseniigiven above. An underwater photograph of one fromthe Maldives is shown in Fig. 8. The population ofjansenii in the Maldives warrants further study.

The hybrids Thalassoma jansenii x T. quinquevitta-tum (Lay and Bennett) and T. nigrofasciatum x T. quin-quevittatum will be documented by Walsh and Randall.

Acknowledgements Thanks are due Mark A. McGrouther of the Aus-

tralian Museum, David H. Catania from the CaliforniaAcademy of Sciences, and Jef frey T. Williams andSusan Jewett of the National Museum of Natural His-tory for the loans of specimens of Thalassoma.Loreen R. O’Hara took X-rays of type specimens of T. nigrofasciatum.

ReferencesAllen, G. R. 1995. Thalassoma robertsoni , a new

species of wrasse (Labridae) from Clipperton Islandtropical eastern Pacific. Revue Française d’Aquari-ologie, 22 (3-4): 75-82.

Bleeker, P . 1856. Beschrijvingen van nieuwe enweinig bekende vischsoorten van Manado enMakassar. Acta Societatis Scientiarum Indo-Neerlandicae, 1:1-80.

Bleeker, P . 1862. Atlas ichthyologique des Indes Orientales Néêrlandaises, publie sous les auspicesdu G ouvernement c olonial n éêrlandais. Tome I.

Scaroïdes et Labroïdes. Frédéric Muller, Amster-dam. xxi + 168 pp.

Cuvier, G. & A. V alenciennes 1839. HistoireNaturelle des Poissons . V ol. 13. Chez Pitois-Levrault, Paris. xix + 505 pp.

Eichler, D. & R. F. Myers. 1997. Korallenfische Zen-traler Indopazifik. Jahr Verlag, Hamburg. 489 pp.

Francis, M. P . 1991. Additions to the fish faunas ofLord Howe, Norfolk, and Kermadec Islands, south-west Pacific Ocean. Pacific Science, 45 (2): 204-220.

Francis, M. P. 1993. Checklist of the coastal fishes ofLord Howe, Norfolk, and Kermadec Islands, south-west Pacific Ocean. Pacific Science, 47 (2): 136-170.

Fricke, R. 1999. Fishes of the Mascarene Islands(Réunion, Mauritius, Rodriguez). An annotated check-list with descriptions of new species . Koeltz ScientificBooks, Koenigstein, Germany. viii + 759 pp.

Heiser, J. B. 1981. Review of the labrid genus Thalassoma (Pisces: Teleostei). PhD thesis. vi + 252pp. Cornell University, Ithaca, NY.

Laboute, P. & R. Grandperrin. 2000. Poissons deNouvelle-Calédonie. Editions Catherine Ledru,Nouméa, 520 pp.

Paepke, H.-J. 1999. Bloch’s Fish Collection in theMuseum für Naturkunde der Humboldt Universitätzu Berlin . A.R.G. Gantner , Ruggell/Liechtenstein.216 pp.

Randall, J. E. 1995. On the validity of the easternPacific labrid fishes Thalassoma grammaticumGilbert and T. virens Gilbert. Bulletin of Marine Science, 56 (2): 670-675.

Randall, J. E., G. R. Allen, & R. C. Steene. 1990.Fishes of the Great Barrier Reef and Coral Sea.University of Hawaii Press, Honolulu. xx + 507 pp.

Randall, J. E. & A. Edwards. 1984. A new labrid fishof the genus Thalassoma from the Pitcairn Group,with a review of related Indo-Pacific species. Journalof Aquariculture and Aquatic Sciences, 4 (2): 13-32.

Randall, J. E. & J. K. L. Mee. 1994. A new labrid fishof the genus Thalassoma from Oman. Fauna ofSaudi Arabia, 14: 302-308.

Randall, J. E. & D. G. Smith. 2001. Thalassoma mas-carenum Fricke, 1999 (Perciformes: Labridae), a syn-onym of T. genivittatum (Valenciennes, 1839).Journal of South Asian Natural History, 5 (2): 117-120.

ERRATUM

aqua 5(4): Fifty new records of shore fishes from theSociety Islands and Tuamotu Archipelago by JohnE. Randall, Philippe Bachet, Richard Winterbottomand Louise Wrobel.A photograph of the goby Paragobiodon lacunicoluswas mistakenly used for the goby Gobiopsis exiguaion p. 161. The correct photo taken by Richard Winterbottom is shown here to the right.

Thalassoma nigrofasciatum, a new species of labrid fish from the south-west PacificThalassoma nigrofasciatum, a new species of labrid fish from the south-west Pacific

Gobiopsis exigua

aqua vol. 7 no. 1 - 2003 8

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KeywordsIchthyology, systematics, Gobiidae, Trimma, new

species, western Indian Ocean

AbstractA new species of the genus Trimma is described.

Trimma volcana is characterized by having largespots on the head and body , moderate to well-devel-oped interorbital and postorbital trenches, scales pre-sent on the pectoral and pelvic fin bases but not onthe cheeks, opercles, or midline of the nape, a poste-rior nostril which is adnate to the eye, and a fifth pelvicfin ray that is 80-90% of the fourth and is brancheddichotomously twice. Trimma volcana has been foundoff Tanzania, Mozambique and the Comores Islands.

ZusammenfassungEine neue Art aus der Gattung Trimma wird be-

schrieben. Trimma volcana kennzeichnet sich durchdie Anwesenheit dunkler Punkte an Kopf und Körper ,mäßig- bis gut entwickelten interorbitalen und postor-bitalen Furchen, Schuppen auf den Basen der Brust-und Bauchflossen aber nicht auf den Wangen, Kiemen-deckeln oder der Mittellinie des Nackens, einem hin-teren Nasenloch das mit dem Auge verwachsen ist undeinem fünften Bauchflossenstrahl mit einer Länge von80 bis 90% des vierten Strahls und der zweimal dicho-tom vergabelt ist, aus. Trimma volcana ist vor Tansania,Mozambique und den Komoren gefunden worden.

RésuméUne espèce nouvelle du genre Trimma est décrite.

Trimma volcana se caractérise par de grosses tâchessur la tête et le corps, des sillons inter- et postorbitauxmodérément à bien développés, des écailles à la basedes pectorales et des pelviennes, mais absentes surles joues, les opercules ou sur la ligne centrale de lanuque, une narine postérieure adnée à l'oeil et, à lapelvienne, un cinquième rayon qui ne fait que 80-90 %

du quatrième et se divise dichotomiquement deux fois.Trimma volcana a été trouvé au large de la Tanzanie,du Mozambique et des îles Comores.

SommarioViene descritta una nuova specie del genere Trimma.

Trimma volcana è caratterizzata dall’avere grandi mac-chie sulla testa e sul corpo, solchi interorbitale e postor-bitale da moderatamente a ben sviluppati, scaglie allabase delle pinne pettorali e pelviche, ma non su guance,opercolo e linea mediana della nuca, narice posterioreadesa al margine dell’occhio e quinto raggio pelvico pariall’80-90% del quarto e con doppia ramificazione dico-tomica. Trimma volcana è dif fusa al largo delle costedella Tanzania, del Mozambico e delle Isole Comore.

IntroductionTrimma are small (to 30 mm SL), usually colourful,

coral reef fishes which can be recognized by the lackof cephalic sensory canal pores, much reducedcephalic sensory papillae pattern, wide gill openingextending to below the vertical limb of the preopercleor anterior to this, lack of spicules on the outer gillrakers of the first gill arch, less than 12 dorsal andanal fin rays, and a fifth pelvic fin ray that is equal toor more than 40% the length of the fourth pelvic finray. There are 36 valid species of Trimma and approx-imately 45 additional species that have yet to bedescribed.

Methods and materialsMethods follow Winterbottom (2002), except pec-

toral and pelvic fin ray branching is described frompreserved material stained with a cyanine blue solu-tion as outlined in Saruwatari et al. (1997). Values forholotype are in bold where appropriate. The speci-mens examined are deposited in the Royal OntarioMuseum (ROM) and the South African Institute ofAquatic Biology (RUSI).

aqua vol. 7 no. 1 - 20039


A new species of Trimma (Gobiidae) from the western Indian Ocean

Richard Winterbottom

Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario,M5S 2C6; and Department of Zoology, University of Toronto, Toronto, Ontario, M5S 1A1.

Email: [email protected]


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Trimma volcana n. sp.Volcano Pygmy Goby (Figs. 1-3)

Holotype: ROM 73474, 17.1 mm SL, female,Comores; Mayotte, Pomanji Is., collected off the northend of airport runway (12º48’12”S, 45º16’18”E), in0.5-5 m by R. Winterbottom and others, 12 Nov 1988.Paratypes: Comores: ROM 59759, 78 (10.4-19.3),Mayotte, Recife de la Prevoyante, south side east ofred buoy (Precaution Reef), lagoon on east side ofMayotte (12º41’30”S, 44º10’00”E), 15-20 m, R. Win-terbottom and co-collectors, 10 Nov . 1988; ROM59760, 2 (18.0-18.3), Mayotte, 100 m north of isleGombe-Doume (12º44’36”S, 45º13’30”E), 3-5 m, R.Winterbottom and co-collectors, 1 1 Nov 1988; ROM59761, 29 (9.5-17.0), Mayotte, north side of isleMalandzamiayatsini (12º40’23.5”S, 45º04’28”E), 6-18m, R. Winterbottom and co-collectors., 15 Nov 1988;ROM 59762, 9 (10.1-14.8), Mayotte, north edge ofBarc Boa Sadia (12º40’22”S, 45º02’17”E), 15-24 m,R. Winterbottom and co-collectors, 17 Nov 1988;ROM 59763, 1 (19.2), collected with the holotype;ROM 1407CS, 7 (12.4-17.9), same as ROM 59759;RUSI 31038, 1 (18.0), Moheli, collected by P . C.Heemstra and co-collectors, 22 Oct 1986.Non-types: Tanzania: RUSI 658, Zanzibar Island.Mozambique: RUSI 898, Wamizi Island.

DiagnosisA species of Trimma with spots on the head and body,

an immaculate cheek, scales on the pectoral and pelvicfin bases but none on the cheek, opercle or midline ofthe nape, an elongate second dorsal spine, interorbitaland postorbital trenches, no scales in the predorsal, anda posterior nasal opening which is adnate to the eye.

DescriptionThe description is based on the holotype and up to

33 specimens. Dorsal fins VI I 10-11, (11 once, n =33), second spine elongate reaching posteriorly to themiddle of the second dorsal fin, the first ray of the sec-ond dorsal fin may be branched or unbranched, allremaining rays are branched; anal fin I 9-11, (x = 9.4, n = 31, 1 1 once), first ray of the anal fin may bebranched or unbranched, all remaining rays arebranched; P 17-18-20, (x = 18.9, n = 28, 17 once),reaching posteriorly to a vertical in line with the ante-rior elements of the anal fin, uppermost one to sevenrays and ventralmost eight to ten rays unbranched,with three to ten branched rays in between; V1 5, nofrenum, basal membrane highly variable, between 10-60-100%, first four rays with 2-3 sequential branches,fifth ray branching dichotomously twice, 80-90% thelength of the fourth ray, fourth ray reaching posteriorlyto the anterior elements of the anal fin. Lateral scales21-24-25 (x = 23.2, n = 29); anterior transverse scales6-7-8 (x = 7.2, n = 30); posterior transverse scales 6-7-8 (x = 7.1, n = 30); opercle, midline of nape andcheek scaleless; 4-5 rows of cycloid scales present onthe pectoral base; cycloid scales present on pelvicbase. Teeth in both jaws consist of an outer row ofcurved, enlarged canines and inner rows of smallerconiform teeth. Tongue is truncate or rounded. Gillopening extends anteroventrally to below posteriormargin of pupil; gill rakers on first arch 2-4 (2 twice) +11-14-15 (11, 13, 14, 15, once) = 14-17, (x = 15.9, n = 19). Anterior nasal opening is a long narrow tube,posterior nasal opening a large rimmed pore adnate toeye. Bony interorbital 1/3 pupil width, with a moderateto well developed interorbital and a slight to well devel-oped postorbital trench. Sensory papillae as in Fig. 2.

Colour when freshly dead ( from two colour slidesfrom Comores): Background colour brown with spotson head and body. Spots on head and dorsal portionof the anterior half of body are yellowish, 1/4-1/3 pupilsized and irregular . Spots on the rest of the body

Fig. 1. Left lateral view of a freshly collected specimen of Trimma volcana (13.6 mm SL female paratype, Mayotte,Comores, ROM 59762). Photo by R. Winterbottom.

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aqua vol. 7 no. 1 - 200311

Richard Winterbottom

grade from yellowish to orange on the mid-portion andfrom orange to red on the caudal peduncle. Spots are1/3 pupil sized and vary in shape (round, oval or irreg-ular). Second dorsal, anal and caudal fins are brownwith orange spots at the base grading to yellow at thedistal portion of the fin. The basal half of the first dor-sal fin is brown, with either yellow or orange spots,and the distal half is hyaline. Paired fins hyaline.Background colour of cheek, opercle and ventral por-tion of head is reddish rather than brown. No spotspresent on cheek.

Colour in alcohol: Background colour straw yellow ,sprinkled heavily with melanophores on the head andchromatophores on the body . Light spots (with little orno pigment) present on the nape, opercle, body andmedial fins. Spots on the head are about 1/4-1/3 pupilsized, quite round, distinct and surrounded by

melanophores. Spots on the body are about 1/3 pupilsized, oval, not clearly defined and surrounded by chro-matophores. Cheek dusky , with no spots. Paired finshyaline with a sprinkling of melanophores at the bases.

EtymologyFrom the Italian, derived from the Latin, Volcanus,

Roman god of fire, in reference to the amber-redspots on an earthy brown background that resemblenumerous small, volcanic vents on a lava field, and issuggestive of the hot-spot origins of the ComoresIslands. To be treated as a noun in apposition.

DistributionTrimma volcana has been found of f the Comores

Islands, Tanzania and Mozambique at depthsbetween 0.5-24 m.

Fig. 3. Left lateral view of Trimma volcana (preserved, 17.4 mm SL female, paratype, Mayotte, Comores, ROM 59759).Photo by R. Winterbottom.

Figs. 2 A-B. Left lateral ( A) and dorsal ( B) views of the head of Trimma volcana (preserved, 17.4 mm SL female,paratype, Mayotte, Comores, ROM 59759). Photos by R. Winterbottom.

A B

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aqua vol. 7 no. 1 - 2003 12


AffinitiesThere are six species of Trimma, which possess

numerous spots on the head and body and a nakednape. In Trimma avidori and T. filamentosus spots arepresent on the centre of each body scale and this cre-ates distinct and even rows of small spots along thelength of the body . The body spots on T. volcana, T.macrophthalma, T . w outsi and T. flammeum arelarger, less numerous and more randomly placed onthe body . However the last three species all havespots on the cheek as does T. filamentosus. Trimmawoutsi, T. macrophthalma and T. flammeum also haveno postorbital trench and the posterior nasal openingis not adnate to the eye. Trimma volcana has no spotson the cheek, a moderately-to well-developed postor-bital trench and a posterior nasal opening that isadnate to the eye.

DiscussionTrimma volcana shares many meristic and morpho-

metric characteristics with T. corallinum, particularlythe well-developed interorbital and postorbitaltrenches and the position of the posterior nasal open-ing (adnate to the eye). Trimma volcana is also verysimilar in appearance to T. flammeum with regard tosize and pattern of spots, the presence of scales onthe pectoral and pelvic bases and elongate dorsalspines. Consequently T. volcana has been previouslymisidentified as both T. corallinum and T. flammeum.

Trimma volcana also appears to form harems. Onelarge lot of specimens from the Comores consistedoverwhelmingly of females. Out of 29 specimens, 23were females and one male (in five specimens thesex could not be determined). In a second lot (78specimens), nine specimens had no discernablepapilla (size range 10.5-12.5 mm SL), 34 specimenshad a papilla but the sex could not be determined withany confidence (10.4-12.5 mm SL), 29 specimenswere female (12.8-19.3mm SL) and six specimenswere questionably male (1 1.0-13.6). At least one ofthe larger females in the second lot was gravid.

AcknowledgementsIt is my pleasure to thank my research assistant,

Margaret Zur, for the data gathering and analysis pre-sented in this paper . Many thanks, too, to my com-panions and co-collectors of the extensive ROMComores material (Peter Benjamin, Colin Buxton,Wouter Holleman, Kathy Kay , Tony Marnewick andRobin Stobbs. Also much appreciated is Tony Kay’svery generous charter rate for his ketch, Before TheWind, which allowed us freedom of movement duringthe 1988 Comores Expedition of the ROM and theSouth African Institute of Aquatic Biology (RUSI).Fieldwork was facilitated through grants from theROM Foundation and a National Sciences and Engi-neering Research Council of Canada Grant (OGP0007619) to the author . The paper published herein

represents Contribution No. 2 of the Centre for Biodi-versity and Conservation Biology of the Royal OntarioMuseum (ROM) to the biological sciences.

ReferencesSaruwatari, T., Lopez, J. A., & Pietsch, T. W. 1997.

Cyanine blue: a versatile and harmless stain forspecimen observation. Copeia, 1997 (4): 840-41.

Winterbottom, R. 2002. Two new species of Trimma(Gobiidae) from the central, western and southPacific. aqua, Journal of Ichthyology and AquaticBiology, 5 (2), 45-52.

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aqua vol. 7 no. 1 - 200313

KeywordsTaxonomy, marine fishes, south-western Pacific,

Gobiidae, Trimma, new species

AbstractA new species of Trimma caesiura species complex,

T. lantana, is described from the north-eastern marginof the Australasian plate and the Solomon Islands, andfour other species of the complex are redescribed. Thisspecies complex is defined by the possession of adeep, steep-sided trench between and posterodorsal tothe orbits. The new species dif fers from its congenersin the presence of two brick-red or brown, dark-edged,rounded blotches over the vertical limb of the preoper-cle, and smaller but similar blotches on the cheekbelow the eye and on the dorsal surface of the snout. Itdiffers further from T. caesiura in having an elongatespine in the first dorsal fin and ventrolateral white spotson the caudal peduncle, and in lacking thin white barson the cheek. Trimma naudei has a club-like red barover the vertical limb of the preopercle and a distinctdark bar over the bases of the pectoral fin rays (the lat-ter diffuse or absent in T. lantana). Two other nominalspecies that appear to belong to this complex areredescribed here. Both T. mendelssohni and T. winter-bottomi have more than a single branch in the fifthpelvic fin ray, usually more pectoral fin rays (a mean of18 vs. 16), and a posterior nasal opening adnate to theanterior margin of the eye (vs. distinctly separate fromthe eye margin). The taxonomic status of two otherspecies in this complex, T. corallinum and T. omanen-sis, is currently under review elsewhere, and thesespecies are not considered further here.

ZusammenfassungTrimma lantana, eine neue Art aus dem Trimma cae-

siura-Artenkomplex, wird von der nordöstlichen Kanteder Australasischen Platte und den Salomon Inselnbeschrieben, und vier weitere Arten aus diesem Kom-plex werden hier neu beschrieben. Dieser Artenkom-plex wird durch die Anwesenheit eines tiefen, steilseiti-gen Grabens zwischen und posterodorsal zu denAugenhöhlen gekennzeichnet. Die neue Art unter-

scheidet sich von ihren Gattungsverwandten durch dieAnwesenheit von zwei ziegelroten oder braunen,dunkel umrandeten Flecken über dem vertikalen Glieddes Vorkiemendeckels, sowie durch kleinere aber ähn-liche Flecken auf der W ange unter dem Auge und aufder dorsalen Oberfläche des Mauls. W eiterhin unter-scheidet sich die neue Art auch von T. caesiura in demsie einen verlängerten Stachel in der ersten Rücken-flosse hat sowie einen ventro-lateralen weißen Punktauf dem Schwanzstiel zeigt; auch sind die schmalenweißen Streifen auf der W ange abwesend. Trimmanaudei hat einen keulenartigen, roten Streifen überdem vertikalen Glied des Vorkiemendeckels und einendeutlichen dunklen Streifen über den Basen der Brust-flossenstrahlen (in T. lantana sind diese Streifen ver-wischt oder ganz abwesend). Zwei weitere nominaleArten, die anscheinend auch zu diesem Komplexgehören, sind hier ebenfalls neu beschrieben worden.Beide Arten, T. mendelssohni und T. winterbottomihaben jeweils mehr als einen einfachen Zweig im fün-ften Bauchflossenstrahl, gewöhnlich mehr Brust-flossenstrahlen (im Durchschnitt 18 im V ergleich zu16), und eine hintere Nasenöf fnung, die mit demvorderen Rand des Auges verwachsen ist (als vergle-ichsweise, deutlich vom Auge abgetrennt zu sein). Diesystematischen Positionen der anderen beiden Artendieses Komplexes, T. corallinum und T. omanensis,werden gegenwärtig anderweitig überprüft, und sindhier somit nicht weiter in Betracht gezogen worden.

RésuméOn décrit Trimma lantana, une nouvelle espèce du

complexe T. caesiura, du bord nord-est de la plaqueaustralasienne et des îles Salomon, et on redécrit qua-tre autres espèces du complexe. Le complexe se car-actérise par la présence d'un profond sillon aux bordsescarpés entre les orbites et en position postérodor-sale. La nouvelle espèce se distingue de ses con-génères par la présence de deux taches arrondies,rouge brique ou brunes, aux bords foncés, sur le limbevertical du préopercule et des taches similaires, maisplus petites, sur la joue, sous l'oeil et sur la face dorsaledu museau. Elle dif fère encore de T. caesiura par une


A new species of the Trimma caesiura species complex (Teleostei: Gobiidae)from the north–eastern margin of the Australian Plate, with a redescription

of the other nominal species in the complex

Richard Winterbottom and Cesar A. Villa

Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, 100 Queen’s Park, Toronto, ON, Canada, M5S 2C6.


7(1)Imp/ok 16-06-2003 11:33 Pagina 13

aqua vol. 7 no. 1 - 2003 14

A n. sp. of the Trimma caesiura complex (Teleostei: Gobiidae), with a redescription of the other nominal species in the complex

épine prolongée dans la première dorsale et par destaches blanches ventrolatérales sur le pédoncule cau-dal et par l'absence de fines barres blanches sur lajoue. Trimma naudei présente une barre rouge enforme de club de golf sur le limbe vertical du préoper-cule et une barre foncée caractéristique sur la base desrayons des pectorales (celle-ci diffuse ou absente chezT. lantana). Deux autres espèces nominales qui sem-blent appartenir à ce complexe sont redécrites ici. T.mendelssohni et T. winterbottomi ont tous les deux plusqu'une branche unique dans le cinquième rayon de lapelvienne, généralement plus de rayons aux pec-torales (une moyenne de 18 contre 16) et une ouver-ture nasale postérieure adnée au bord antérieur del'oeil (alors que nettement séparée du bord de l'oeilchez les autres). Le statut taxonomique de deux autresespèces de ce complexe, T. corallinum et T. omanen-sis, est actuellement à l'étude ailleurs et ces espècesne sont pas davantage prises en compte ici.

SommarioUna nuova specie del complesso Trimma caesiura, T.

lontana, viene descritta sulla base di esemplari raccoltilungo il margine nordorientale della piattaforma aus-tralasica e le Isole Solomone e altre quattro specie delcomplesso vengono ridescritte. Questo complesso dispecie è definito dal possedere un profondo solco nellaregione interorbitale posterodorsale. La nuova speciesi distingue dai suoi congeneri per la presenza di duemacchie tondeggianti color rosso mattone o brunastrosopra il ramo verticale del preopercolo e per la pre-senza di altre macchie simili ma più piccole sulleguance al di sotto dell’occhio e sulla superficie dorsaledel muso. Si distingue inoltre da T. caesiura per avereuna spina più allungata nella prima pinna dorsale e unamacchia bianca ventrolaterale sul peduncolo caudale eper l’assenza di sottili strie bianche sulle guance.Trimma naudei ha una barra rossa a forma di clavasull’asse verticale del preopercolo e una distinta barrascura alla base dei raggi della pinna pettorale (quest’ul-tima diffusa o assente in T. lontana). Due altre specienominali che sembrano appartenere a questo comp-lesso sono ridescritte. Sia T. mendelssohni che T. win-terbottomi hanno più di una ramificazione nel quintoraggio pelvico, generalmente un maggior numero diraggi pettorali (mediamente 18 vs. 16) e l’aperturanasale posteriore contigua con il margine anteriore del-l’occhio (vs. distinta dal margine oculare). Lo stato tas-sonomico di altre due specie del complesso, T.corallinum e T. omanensis, è attualmente in corso direvisione e non viene considerato in questo lavoro.

IntroductionTrimma is a speciose genus of small (< 30 mm SL)

Indo-Pacific coral reef gobiid fishes. Within the taxon,certain apparently monophyletic sub-units are apparent.The five species described here seem to belong to sucha unit, based on the shared possession of apparently

specialized trenches between and behind the orbits,and a triangular shaped ridge (apex pointing anteriorly)formed by the frontal bones marking the posterior mar-gin of the trench system. The trench system is thusessentially a Y-shaped trough, with the stem of the Ydirected anteriorly (see Fig. 1). Such a configuration isnot known in any other nominal Trimma species, withthe exception of Trimma corallinum and Trimma oma-nensis, whose current relationship has yet to beresolved, and are therefore excluded. The absence ofthe trench system in most other Trimma species, and inpotential outgroups such as Trimmatom and Priolepis(Winterbottom and Burridge, 1992), is hypothesized torepresent a pleisiomophic condition, and our nullhypothesis is thus that the trenches represent a derivedcondition. The species complex as described here maynot form an exclusive monophyletic group, as a surveyof the genus has revealed approximately 10 unde-scribed species that appear to possess such trenches,or a configuration of the frontal region analogous tothem. Our purpose is to describe a new species fromthe south-eastern part of the Indonesian triangle andAustralia, as well as to provide a redescription and keyto the nominal species treated here. This species com-plex can be artificially divided into two groups based onsimilarity of pectoral fin ray count, fifth pelvic fin raybranching pattern, and posterior nasal opening positionrelative to the eye. Trimma caesiura, T. naudei, and thenew species form one group based on a single dichoto-mous branch in the fifth pelvic fin ray , a pectoral fin raycount averaging 16 but not exceeding 18, and a poste-rior nasal opening separated by about half a pupil widthfrom the anterior margin of the orbit. Trimma winterbot-tomi and T. mendelssohni form the other group, in whichthere is more than one type of branching pattern in thefifth pelvic fin ray, an average of 18 pectoral fin rays (atotal of 23 rays in some cases), and a posterior nasalopening which is adnate to the anterior margin of theorbit.

Materials and methodsMethods follow Hubbs and Lagler (1958) with the

exception that lateral scales are counted from the scaleoverlapping the distal end of the hypural plate anteriorlyalong the midlateral septum to the axil of the pectoral fin,and transverse scale row counts begin at the scale adja-cent to the anus and follow the scale rows anterodor-sally and posterodorsally to the scale adjacent to thebase of the dorsal fins. Values for holotype of the newspecies are in bold where appropriate. Abbreviations forrepositories of the material examined are as follows:ANSP – Academy of Natural Sciences of Philadelphia;BPBM – Bernice P. Bishop Museum, Honolulu; ROM –Royal Ontario Museum, Toronto (note: entries contain-ing the letters ‘CS’denote cleared and stained material);USNM – National Museum of Natural History, Washing-ton (formerly United States National Museum); W AM –Western Australian Museum.

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Figs. 1 A-E. Interorbital trenches in: A - Trimma caesiura, 25.5 mm SL male, Fiji. ROM 46085; B - T. lantana, 20.8 mmSL female (holotype), Solomon Is., ROM 46039; C - T. winterbottomi, 20.5 mm SL male, Thailand, ROM 68103;D - T. naudei, 26.5 mm SL female, Comores, ROM 59796; E - T. mendelssohni, 20.9 mm SLmale, Comores, ROM 59791.Photos by R. Winterbottom.

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Artificial key to the nominal species of Trimma with trenches

1. One dichotomous branch in the 5th pelvic fin ray; anaverage of 16 (no individuals exceeding 18) pectoralfin rays; a posterior nasal opening separate from theanterior margin of the eye by about half the width ofthe pupil ...................................................................2

Multiple dichotomous or sequential branches in the 5thpelvic fin ray; an average of 18 or more pectoral finrays; a posterior nasal opening adnate to the ante-rior margin of the eye ..............................................4

2. The second spine of the first dorsal elongate; a dis-tinct, white ventro lateral spot pattern; a spotted orpatchy colour pattern over the vertical limb of thepreopercle ................................................................3

The second spine of the first dorsal not elongate; noventrolateral white spots; two thin, diffuse, whitelines below the eye, and one over the vertical limb ofthe preopercle (Western Pacific)....Trimma caesiura

3. No spots, bars or blotches below the eye or on dor-sal surface of snout; a uniform dusky orange / greycolour below the eye extending anteriorly to thesnout, with orange / red blotches against a white /grey background on the head and nape, and a sim-ilar orange / red ground colour on the trunk withwhite stripes and spots; a vertical bar on the poste-rior margin of the pectoral base formed bymelanophores; a single, large, dorsally-expanded,orange blotch with dark edging (resembling a club)over the vertical limb of the preopercle (W esternIndo-Pacific).......................................Trimma naudei

A single red / orange bar followed by a series of twoirregular blotches below the eye; the head and dor-sal surface of the snout with red / orange blotchesagainst an of f-white / grey background anteriorly ,darkening towards the snout where it is grey / blue(steel blue live), and with a red / orange trunk colourfading to yellow caudally with white stripes andspots; a slightly darkened posterior margin of thepectoral base (preserved specimens); two distinctblotches, one under the other vertically, over the ver-tical limb of the preopercle, both edged withmelanophores (not resembling a club) (AustralianPlate and Solomon Islands) .............Trimma lantana

4. Second dorsal fin with 9-10 elements; multiplebranches in the 5th pelvic fin ray; scales present onthe head, opercle, pectoral base and prepelvicregion; two small, fleshy lappets on the dorsal sur-face of the nape (bright white in life); no groups ofmelanophores above the posterior margin of theopercle; an overall mottled brown / red body colourwith five white, diffuse bars along the dorsum (West-ern Indian Ocean) ..................Trimma mendelssohni

Second dorsal fin with 10-1 1 elements; two sequentialbranches in the 5th pelvic fin ray (occasionally onedichotomous branch); no scales present on thehead, opercle, pectoral base or prepelvic region; no

fleshy lappets on the dorsal surface of the nape; atriangular group of melanophores above the dorsalmargin of the opercle; an overall orange / red bodycolour with five poorly-defined white, elongatedspots along the dorsum (Persian Gulf – Thailand) ....... ............................................Trimma winterbottomi

Trimma lantana n. sp.(Figs. 1B, 2, 3, 13)

Trimma sp. Burgess et al., 1990: Plate 492, 577; Allen,1997: Plate 90, 12; Randall 1998: 251, Fig. 76;Winterbottom, 1984 (brief comparison with T. naudei).

Trimma sp. 1 Kuiter 1992: 236, Fig. B.

Material Examined Thirty-three specimens examined, 1 1.0 – 29.4 mm

SL, all collected with quinaldine or rotenone.

Holotype: Honiara, 2 km west of Point Cruz, Guadal-canal, Solomon Islands, Nichols and Evans, 20.8 mmSL, ROM 46039, female.Paratypes: Ashmore Reef, Timor Sea, north-easternAustralia, Allen et al., 5 (18.7-22.9), WAM P29047-034;2 (19.4-26.5), WAM P29051-024; Escape Reef, GreatBarrier Reef, Queensland, Australia, Paxton et al., 1(19.8), ROM 40611; Escape Reef North, Great BarrierReef, Queensland, Australia, A yling et al., 1 (21.1),ROM 40539; Endeavor Island, Great Barrier Reef,Queensland, Australia, Tyler and Smith, 1 (11.0),USNM 295142; Lizard Island, Great Barrier Reef,Queensland, Australia, Winterbottom e t al ., 4 (18.1-21.3), ROM 39328; with 2 specimens removed forclearing and staining, (18.6-18.7), ROM 1172CS; TijouReef, Great Barrier Reef, Queensland, Australia,AMS/AIMS party , 3 (21.0-21.9), ROM 40551; YongeReef, Great Barrier Reef, Queensland, Australia, Win-terbottom et al., 1 (19.2), ROM 39333; Pt. Quobba,north-western Australia, Hutchins et al., 5 (22.1-26.9),WAM P27920-037; Kranket Island, Papua NewGuinea, Randall, 1 (15.4), BPBM 32680; Manus Island,Papua New Guinea, Allen and Knight, 1 (12.5), W AMP27827-046; Honiara, Guadalcanal, Solomon Islands,Nichols 2 (18.9-20.6), ROM 46044; collected with holo-type, 5 (11.5-29.4), ROM 73436.

DiagnosisA species of Trimma with a frontal ridge behind the

orbits that slopes steeply into a wide interorbital trenchanteromedially, and into postorbital trenches laterally(see description below). Posterior nasal opening sepa-rate from anterior margin of eye; second spine of firstdorsal elongate, third spine not elongate; fifth pelvic finray branched once dichotomously; body colour of f-white / grey anteriorly with grey / blue snout, red /orange trunk grading to yellow caudally, with five whitedorsolateral spots between peduncle and origin of first

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dorsal, and two white ventrolateral spots between thepeduncle and anal fin; a red / orange bar and two irreg-ular blotches under orbit; slightly darkened posteriormargin of pectoral base.

DescriptionThe description is based on 33 specimens, 11 males,

19 females, and 3 juveniles (including 2 cleared andstained).

Dorsal fins VI + I 7-8-9 (x = 8.0), second spine of D 1elongate in both sexes, extending posteriorly betweenorigin of second ray of D 2 to mid-peduncle whenadpressed, third spine usually not elongate; anal fin I 7-8-9 (x = 8.4); pectoral fin rays 15- 16-18 (x = 16.4),uppermost three and lowermost five rays unbranched,with about nine branched rays in between; pelvic fin I 5,no fraenum, basal membrane vestigial; fifth pelvic fin raybranched once dichotomously, 50-70-90% (x = 71.0%)length of fourth ray. Lateral scales 21-24-25 (x = 23.6),anterior transverse scales 6- 7-9 (x = 7.1), posteriortransverse scales 6-8 (x = 7.2), predorsal scale rows 5-7-8 (x = 6.6); scales on opercle, pectoral base, and pre-pelvic region cycloid; all other body scales ctenoid; 1-2(x = 1.2) scales on anterodorsal corner of opercle; 1-3(x = 1.7) scale rows on pectoral base; 3- 4-7 (x = 4.4)scale rows in prepelvic region; specialized triangularscale with elongate apex pointing posteriorly on thepelvic base (between innermost pelvic fin rays). Teeth inboth upper and lower jaw with large, curved, spaced,canines in outer row, with smaller irregular inner rows ofconical teeth; innermost row of lower jaw with spaced,somewhat enlarged, canines less than half the size ofouter; tongue shape rounded or bilobed. Gill openingextending to below mid-pupil; gill rakers on outer surfaceof first arch 3-4 + 15-16-17 (n = 6, x = 3.4 + 15.7). Ante-rior nasal opening tubular and slightly flared; posteriornasal opening a pore with raised rim, its posterior mar-gin about half a pupil width from anterior margin of eye.

Trenches: the frontal bones form a ridge nearly verti-

cal to the horizontal plane, behind the posterior marginsof the orbits. Trimma lantana has a frontal ridge with anoverall triangular shape in dorsal profile (apex pointinganteriorly, and slightly concave at the tip) (see Fig. 1B).The apex of the ridge slopes steeply anteroventrally intoa trench between the orbits, in the frontal midline.Behind the orbits, the ridge ends abruptly (vertical drop-off) forming the posterior wall of the U-shaped postor-bital trenches. The anterior walls of the postorbitaltrenches, and the lateral walls of the interorbital trench,are formed by the vertically sided rims of the bony mar-gin of the frontal bones. The postorbital trenches beginfrom the posterior margin of the orbits, continueanterodorsally around them, and merge in the interor-bital region to form the interorbital trench, which endsbefore the snout.

Colour in life (based on underwater photos by G.Barrall and G. R. Allen, and freshly-collected speci-mens, based on colour slides, from the Great BarrierReef, Australia, and Guadalcanal, Solomon Islands) :the anterior half of body is of f-white / grey , darkeningtowards the snout where it is steel blue (live) or grey /blue (freshly collected), with numerous, irregular red /orange blotches margined with melanophores on thedorsal surface of the snout, head and cheek (about halfa pupil width on cheek). The area below the orbit has asingle complete red / orange bar under the anteriorhalf, followed by two vertically-aligned, round blotches,the dorsal most half a pupil width, the other pupil width,under the posterior half. There are two vertically-aligned, separated, irregular red / orange blotches mar-gined with melanophores on the vertical limb of the pre-opercle. The blotches gradually become larger, lackmelanophores, and are irregularly-positioned posteri-orly, forming a red / orange ground colour on the trunk,which grades to orange (live) or yellow (freshly-collected) on the peduncle. The anterior off-white / greybackground is reduced to irregular stripes andblotches, in between and around the red / orange

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Fig. 2. Trimma lantana, 19.3 mm SL, freshly-collected, female, Lizard Island, Great Barrier Reef, ROM 39333. Photo byR. Winterbottom.

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blotches, and gradually become smaller posteriorly .The best-developed of f-white / grey stripe is more orless zigzag in shape and passes along the sides of thebelly. Across the dorsum are three similar white spots,approximately 75% of pupil width. The first occurs atthe origin of the first dorsal fin, the second over thebase of the fifth spine of the first dorsal, and the third atthe origin of the second dorsal fin. A fourth white spot,half a pupil width, is situated under the origin of theeighth ray of the second dorsal fin. A fifth white spot,pupil width, is situated across the dorsal margin of themid-peduncle, about an equal distance anterior to thefirst procurrent caudal fin ray (see Fig. 3). Two whitespots occur behind the ventral margin of the anal fin.One spot, 50% pupil width, is situated on the ventralmargin of the mid-peduncle (just posterior to the posi-tion of the dorsal spot). The second white spot, pupilwidth, is situated about a pupil width behind the anal fin.Scale pockets on the body are more intensely pig-mented than their surroundings (see Fig. 2). There is abasal red stripe in the first dorsal fin, which continues inthe second dorsal fin, breaking up into red spots poste-riorly. The second dorsal fin has rows of similar spotsabove the basal stripe, followed by a row of red /orange streaks along the fin elements. The rest of thefirst and second dorsal fins, up to their distal margins,are hyaline. Oblique rows of yellow streaks and spotsmay be present in the caudal fin, which are most promi-nent in the dorsal half of the fin and become irregularlyarranged posteriorly. The anal fin is similar in appear-ance to the second dorsal fin. The pectoral and pelvicfins are hyaline, tinged with red.

Colour in alcohol: the background colour is a palestraw yellow sprinkled lightly with chromatophores pos-teriorly, and more heavily so anteriorly . Three lightbands of chromatophores occur below the eye. Thereare numerous scattered, irregular light spots around thehead and nape, outlined with melanophores, extendingto the middle of the first dorsal fin. There is a faint verti-cal band of melanophores on the posterior margin of thepectoral fin. The triangular perimeter of the frontal ridge

of the head is heavily sprinkled with melanophores. Anaggregate of melanophores, pupil width, may at timesbe present on the dorsum of the mid-peduncle.

Affinities and DiscussionThe new species is most easily confused with T.

naudei and T. caesiura, both of which resemble it inappearance and meristic values. Anteriorly, T. caesiurahas a more conservative red / orange body colorationthan does T. lantana and T. naudei, in which the ante-rior half of the body is composed of irregular orange andred blotches against an off-white / grey (grey / blue inthe case of T. lantana) background. Trimma naudei canmost easily be distinguished from T. lantana by its dark,vertical pectoral bar and an orange club-shaped blotchover the vertical limb of the preopercle. In life, T. lantanahas no pectoral bar, although in preserved specimens afaint group of melanophores may be present along theposterior margin of the pectoral fin base. In addition tothe steel blue tinge to the anterior portion of the head(darkening anteriorly) in life, T. lantana possesses a red/ orange bar and a series of two blotches below the orbitand discreet rounded blotches on the dorsal surface ofthe snout (vs. a more uniform dusky orange / grey washbelow the eye and on the dorsal surface of the snoutwith no distinctive markings in T. naudei). Trimmacaesiura has diffuse white lines on the dusky, faded red/ orange background of the cheek below the eye (andalso lacks distinct markings on the dorsal surface of thesnout), and differs further from the other two species inlacking ventrolateral white markings on the posteriorhalf of the body, no elongate spine in the first dorsal fin,and with a rounded, or convex apex of the frontal ridgein dorsal profile (vs. white spots or stripes present onventrolateral body, an elongate spine in first dorsal fin,and apex of frontal ridge slightly concave).

Specimens of these three species in 70% ethyl alco-hol have a pale straw-coloured body, and spots on thebody that are typically outlined by melanophores.Trimma lantana retains the irregular head and facialpattern in the form of melanophore outlines, but the

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Fig. 3. Trimma lantana. Underwater photograph of live specimen, northern Great Barrier Reef. Photo by G. Allen.

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body spot pattern is not as well-defined as in T. cae-siura, in which the spots may remain as irregular out-lines. Trimma naudei retains the dark pectoral bar, andthe outline of the club-shaped marking on the verticallimb of the preopercle (the interior of which is paler thanthe rest of the body) as well as the dorsolateral andventrolateral spot / stripe patterns which are edged bymelanophores.

Winterbottom (1984), in his redescription of 10 speci-mens of T. naudei from the Chagos Archipelago, pro-vided a brief comparison with T. lantana, saying that “Aspecies somewhat similar to T. naudei occurs on theGreat Barrier Reef of Australia.”(1984:708). He cor-rectly identified some of the distinguishing featureslisted for T. naudei, but erred in stating that the unde-scribed species (= T. lantana) lacked scales on thedorsal margin of the opercle.

The new species has been referred to informally asTrimma RW sp. 56.

EtymologyNamed for the ubiquitous tropical plant, Lantana, the

inflorescences of which are a varying assortment ofyellow, orange, purple, and reddish-pink flowers, inallusion to the colourful nature of the new species andthe rounded blotches on the head. To be treated as anoun in apposition.

DistributionTrimma lantana is currently known from Australia,

Papua New Guinea, and the Solomon Islands. Speci-mens were collected at depths ranging from 3-25 m,within crevices and caves of mostly dead coral on asandy / rubble bottom. See Fig. 13 for a spot distribu-tion map of all species treated in this paper .

Trimma caesiura Jordan and Seale(Figs. 1A, 4, 5, 13)

Trimma caesiura Jordan and Seale 1906: 391 (typelocality: American Samoa)


orbits that slopes steeply into a wide interorbital trenchanteromedially, and into postorbital trenches laterally(see description below). Posterior nasal opening sepa-rate from anterior margin of eye; no elongate first dor-sal spines; fifth pelvic fin ray branched once dichoto-mously; body colour red / orange with four white dorso-lateral spots between caudal region and origin of sec-ond dorsal; no white ventrolateral spots; facial patternwith two thin, dif fuse white diagonal lines under orbitand one over preopercle.

Description The description is based on 22 specimens, 11 males,

9 females and 2 cleared and stained specimens, 13.3-27.0 mm SL.

Great Astrolabe Reef, Fiji, 7 (13.3-22.5), ROM 46080;Viti Levu, Fiji, 6 (22.9-25.3) ROM 46085; Coral Sea,New Caledonia, 4 (1 1.9-27.4), ROM 64326; 3 (18.8-25.4), ROM 64402; Great Astrolabe Reef, Fiji, 2cleared and stained (21.3-22.0), ROM 989CS.

Dorsal fins VI + I 7-8 (x = 7.7), second and third D 1spines not elongate; anal fin I 8-9 (x = 8.1); pectoral finrays 15-19 (x = 16.7), uppermost seven and lowermostfour rays unbranched, with about five branched rays inbetween; pelvic fin I 5, no fraenum, basal membraneusually vestigial (one instance of full membrane); fifthpelvic fin ray branched once dichotomously , 50-90% (x = 64.1%) length of fourth ray . Lateral scales 23-26 (x = 24.6), anterior transverse scales 7-9 (x = 8.1), pos-terior transverse scales 7-9 (x = 7.8), predorsal scalerows 7-9 (x = 7.3); scales on opercle, pectoral base,and prepelvic region cycloid; 1-2 (x = 1.2) scales oftenpresent on anterodorsal corner of opercle (42% ofspecimens); pectoral base with 0-3 (x = 1.3) rowsscales; 3-4 (x = 3.9) scale rows in prepelvic region; spe-cialised triangular scale with elongate apex pointingposteriorly on the pelvic base. Teeth in both upper andlower jaw with large, curved, spaced, canines in outerrow, with smaller irregular inner rows of conical teeth;

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Fig. 4. Trimma caesiura, 23.6 mm SL, freshly-collected, male, New Caledonia, ROM 63917. Photo by R. Winterbottom.

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innermost row of lower jaw with spaced canines greaterthan half the size of outer; tongue rounded to bi-lobed.Gill opening to below the mid-pupil; gill rakers on firstarch 2-3 + 16-17 (n = 7, x = 2.7 + 16.3). Anterior nasalopening tubular and slightly flared; posterior nasalopening a pore with raised rim, about half a pupil widthfrom anterior margin of eye.

Trenches: trenches as in Trimma lantana . Trimmacaesiura has a frontal ridge with an overall triangularshape (apex pointing anteriorly, with convex tip), in dor-sal profile (see Fig. 1A). The apex of the ridge slopessteeply, anteroventrally into a deep trench between theorbits, in the frontal midline. Behind the orbits, the ridgeabruptly ends forming the posterior wall of deep pos-torbital trenches.

Colour in life: (based on underwater photos by H.Nagano, M. Takata, and R. Myers, freshly-collectedspecimens, based on colour slides, from the Coral Sea,New Caledonia, Okinawa, Marshall Islands, Fiji, andGuam): the body pattern consists of a red / orangebackground with four irregular white spots, marginedwith melanophores, on the dorsum, extending from theposterodorsal margin of the peduncle to the origin ofthe second dorsal. A white line zigzags across the dor-sal surface of the head and nape. The white dorsalspots decrease in size posteriorly. The first spot, slightlylarger than pupil width, is situated under the origin ofthe second dorsal, the second, about pupil width,occurs under the origin of the eighth ray of the seconddorsal. A less regular white spot, slightly less than pupilwidth, occurs on the peduncle. The last spot occurs onthe posterodorsal margin of the peduncle. Scale pock-ets on the body are more intensely pigmented abovethe midline. The ventrolateral body pattern typicallylacks white spots, and is composed of scale pocketswith white / silver centres (see Fig. 4). The area underthe orbit is a mix of dusky, faded red /orange colourdusted in melanophores. Two parallel, dif fuse whitelines, angled at 25° posteriorly, about a pupil widthapart, often occur under the orbit. A third, similar line is

situated across the preopercle (see Fig. 5). There is aseries of basal red spots in the first dorsal fin, whichcontinues along the base of the second dorsal fin. Theelements of the first dorsal are tinged red to their tips.The second dorsal fin has rows of similar orange spotsabove the basal spots. The rest of the dorsal fin is hya-line. The anal fin is hyaline, with a basal orange stripe.The pectoral and pelvic fins are hyaline, with a redtinge. The caudal fin has two rows of crescent-shaped,elongated, orange spots.

Colour in alcohol: the body colour is a pale strawyellow with scale pockets margined with melanophor-es. Melanophores also outline four spots on the dor-sum corresponding to the four white spots in live /freshly collected specimens. The base of the pectoralfin is dusky, with no distinct vertical bar.

Affinities and Discussion(See T. lantana).

DistributionTrimma caesiura is currently known from China

(Pratas Reef), Fiji, Guam, Japan, Loyalty Islands, Mar-shall Islands, New Caledonia, Palau, Papua NewGuinea, Samoa, and the Solomon Islands. Specimenswere collected at depths ranging from 3 – 25 m, withincrevices and caves of mostly dead coral on a sandy /rubble bottom.

Trimma naudei Smith(Figs. 1D, 6, 7, 13)

Trimma naudei Smith 1956: 828 (Mahé, Seychelles);Winterbottom, 1984: 708-709 (redescription)

Trimma sp. Akihito et al., 1988: 245 (Ryukyu Islands)


orbits that slopes steeply into a wide interorbital trenchanteromedially, and into postorbital trenches laterally

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Fig. 5. Trimma caesiura, live, Palau. Photo by H. Nagano.

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(see trenches in description below). Posterior nasalopening separate from eye; second spine of first dorsalfin elongate, third spine occasionally elongate; fifthpelvic fin ray branched once dichotomously; bodycolour orange / red with seven dorsolateral white spotsbetween posterodorsal margin of the peduncle and ori-gin of first dorsal, and with four ventrolateral spotsbetween posteroventral margin of peduncle and originof last anal ray; facial pattern uniform dusky orange /grey colour under orbit, with an orange vertical blotch,edged in melanophores on vertical limb of the preoper-cle resembling a club; conspicuous bar of melanophor-es extending vertically across posterior margin of pec-toral base.


10 females and 2 cleared and stained specimens, 17.5-27.2 mm SL. Salomon Atoll, Chagos, 5 (19.2-25.3),ROM 4031 1; Salomon Atoll, Chagos, 4 (17.5-25.0),ROM 40313; South Male Atoll, Maldives, 4 (19.8-24.7),ROM 54948; Moheli, Comores, 4 (22.1-27.2), ROM59796; Mayotte, Comores, 3 (19.1-21.7), ROM 59793;Peros Banhos, Chagos, 2 (21.5-22.0), ROM 739CS.

Dorsal fins VI + I 7-8 (x = 7.4), second D 1 spine elon-gate in males, extending posteriorly between origin ofthird ray of D2 to past peduncle when adpressed; thirdspine extending to origin of second ray of D 2; anal fin I7-9 (x = 8.3); pectoral fin rays 15-17 (x = 15.9), upper-most three and lowermost five rays unbranched, withabout eight branched rays in between; pelvic fin I 5, nofraenum, basal membrane vestigial; fifth pelvic fin raybranched once dichotomously , 60-85 % (x = 70 %)length of fourth ray. Lateral scales 23-26 (x = 24.4),anterior transverse scales 6-9 (x = 7.5), posterior trans-verse scales 6-8 (x = 7.3), predorsal scale rows 6-8 (x = 6.9); scales on opercle, pectoral base, and pre-pelvic region cycloid; all other body scales ctenoid; 1-2 (x = 1.7) scales on anterodorsal corner of the opercle;1-2 (x =1.9) scale rows in pectoral base; prepelvic

region with 3-5 (x = 4.3) scale rows; specialised trian-gular scale with elongate apex pointing posteriorly onthe pelvic base. Teeth in both upper and lower jaw withlarge, curved, spaced, canines in outer row, withsmaller irregular inner rows of conical teeth, and inner-most row of lower jaw with spaced canines less thanhalf the size of outer; tongue rounded to bi-lobed. Gillopening to below the mid pupil; gill rakers on first arch3-4 + 14-16 (n = 20, x = 3.3 + 14.6). Anterior nasalopening tubular and slightly flared; posterior nasalopening a pore with raised rim, about half a pupil widthfrom anterior margin of eye.

Trenches: trenches as in Trimma lantana . Trimmanaudei has a frontal ridge with an overall triangularshape (apex pointing anteriorly, with deep concave tip),in dorsal profile, similar to that of Trimma lantana (seeFig. 1D). In the frontal midline, the apex of the ridgeslopes steeply, anteroventrally into a deep trenchbetween the orbits. Behind the orbits, the ridge abruptlyends forming the posterior wall of deep, narrow postor-bital trenches.

Colour in life (based on underwater photos by Y.Adachi, and J. Randall, and on freshly-collected speci-mens, based on colour slides, from the Phuket, Thai-land; Bolinao, Philippines; Chagos; Comores; andOrchid Island, Taiwan): the body pattern consists of anorange / red background with seven round white spots,margined with melanophores, on the dorsolateral sur-face, extending from the posterodorsal margin of thepeduncle to the origin of the first dorsal. The anteriorhalf of body is white / grey with numerous, irregularorange/red blotches margined with melanophores onthe head and nape. The blotches gradually becomelarger posteriorly, lack melanophores, and become lessdistinct, forming an orange / red ground colour on thetrunk. The anterior white / grey background is reducedto irregular stripes and blotches, in between andaround the coalesced orange / red blotches, and grad-ually become smaller posteriorly . The best developedoff-white stripes are more or less zigzag in shape, and

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Fig. 6. Trimma naudei, 26.7 mm SL, freshly-collected, male, Comores, ROM 59796. Photo by R. Winterbottom.

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pass along the sides of the dorsum and belly . The fol-lowing white spots are all about pupil width in diameter(or slightly larger unless stated otherwise) and sepa-rated from the dorsal midline and each other by aboutthe same distance. The first dorsolateral white spot, issituated at the origin of the first dorsal spine. The sec-ond white spot occurs between the origins of the fifthand sixth spine of the first dorsal fin. A third white spot(about one third of pupil width), occurs at the origin ofthe second dorsal (see Fig. 6). The fourth white spot issituated closer to the midlateral line below the thirdspot, and may become incorporated into the whitestripe zigzagging along the dorsum. The fifth spot is sit-uated at the same distance below the dorsum, from thefourth spot, and of the same diameter as the precedingspots. The sixth white spot is located along the dorsumof the mid-peduncle. The last dorsolateral spot islocated on the posterodorsal corner of the peduncle,and is slightly less than pupil width, although it may attimes form a bar extending to the midlateral region ofthe peduncle (see Fig. 7). The ventrolateral white spotsgradually become smaller and more irregular posteri-orly. The first spot is situated at the origin of the lastanal-fin ray, and may be incorporated into the ventralzigzagging white stripe (see Fig. 6). The second whitespot is located on the mid-peduncle, slightly behind thecorresponding dorsal white spot. The most posteroven-tral white spot is approximately half a pupil width indiameter, and is situated on the posteroventral cornerof the peduncle. The scale pockets on the body aremore intensely pigmented along the midline, and haveorange / red centres in life, or white when freshly-col-lected (see Fig. 6). The pectoral base has a solid verti-cal bar of melanophores set against a white back-ground (see Fig. 6). Occasionally, the pectoral bar isseparated slightly into upper and lower halves (see Fig.7). The area under the orbit is a dusky orange / greycolour, lightly dusted in melanophores, and with noobvious markings. The preopercle has a large vertical

bar, outlined with melanophores and with an orangeinterior, extending from the base of the ascending limbof the preopercle to behind the lateral midpoint of theeye. The dorsal segment of the bar is expanded givingit a club-like appearance (see Fig. 6). There is a basalorange stripe in the first dorsal fin that continues in thesecond dorsal. The second dorsal has another faintstripe, which then breaks up into faint orange spots dis-tally. The first dorsal fin is tinged red to the tips of thespines; the second dorsal fin is hyaline distally . Theanal fin is similar in appearance to the second dorsalfin. Oblique rows of orange / yellow streaks and spotsare present in the caudal fin, becoming irregular dis-tally. The pectoral and pelvic fins are tinged orange withhyaline tips.

Colour in alcohol: the body colour is pale, straw yel-low with scale pockets margined with melanophores.The club-shaped blotch on the preopercle remains asan outline of melanophores with a pale interior . Thepectoral bar is conspicuous in preserved specimens,allowing for easy identification.

Affinities and Discussion(See T. lantana)Examination of additional material of T. naudei did

not support Winterbottom’s 1984 tentative suggestionof sexual dimorphism in eye diameter as a percentageof head length. The specimens agree with Winterbot-tom’s (1984) redescription of T. naudei in all otherrespects.

DistributionTrimma naudei is currently known from the Chagos

Archipelago, China, the Comores, Indonesia, Japan,Malaysia, Maldives, Mauritius, Philippines, Rodriges,Seychelles, Thailand, and Viet Nam. Specimens werecollected at depths ranging from 9-22 m, over coralpatches interspaced with fine sandy bottoms, and mayoccasionally form small schools.

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Fig. 7. Trimma naudei. Underwater photograph of live specimen, Maldives. Photo by J. E. Randall.

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Trimma mendelssohni (Goren)(Figs. 1E, 8, 9, 10, 13)

Quisqulius mendelssohni Goren, 1978:191 (Sinai)Trimma mendelssohni Winterbottom, 1995 (Gulf of

Aqaba, brief diagnosis)

DiagnosisTrimma mendelssohni has a frontal ridge that slopes

variably (steeply, and with a ridge, to gradually , withouta ridge) into the interorbital trench. The postorbitaltrenches may be steep – or shallow-sided (see descrip-tion below). Posterior nasal opening adnate to eye; sec-ond spine of first dorsal fin occasionally elongate; thirdspine not elongate; eight or more elements in the sec-ond dorsal; fifth pelvic fin ray branched multiple timesdichotomously, appearing bushy; 18 or more pectoralrays; scaled head; body colour brown / red with five dif-fuse white vertical bars, between mid-peduncle to justanterior to origin of first dorsal; facial pattern with twowhite bars under orbit, and one irregular white bar oververtical limb of the preopercle; a pair of thin fleshy lap-pets, half a pupil width apart, a quarter pupil width beh-ind the frontal ridge, on either side of dorsal midline ofthe nape; the epibranch of first gill arch lacks gill rakers.

DescriptionTwenty-two specimens examined, 10.4 –24.6 mm SL.

Moheli, Comores, 4 (10.4-16.4), ROM 59791; Gulf ofAqaba, Red Sea, Israel, 3 (10.4-12.7), USNM 295141;Sinai Peninsula, Red Sea, Israel, 3 (17.5-21.7), BPBM18331; St. Brandon’s Shoals, Cargados Carajos, Mau-ritius, 2 (20.9-22.1), USNM 264549; Curieuse Island,Seychelles, 2 (16.5-20.3), USNM 295313; MahéIsland, Seychelles, 2 (16.2-19.3) USNM 295235;Round Island, Seychelles, 4 (17.0-24.6), USNM295143; and 2 cleared and stained, Cargados Carajos,Mauritius, (18.6), ex. USNM 264706; Moheli, Comores,(16.4), ROM 1743CS (ex ROM 59791).

Dorsal fins VI + I 8-9 (x = 8.7), second D 1 spine elon-gate, reaching posteriorly between origin of third ray ofD2 to past peduncle when adpressed, third spine notelongate; anal fin I 7-9 (x = 8.6); pectoral 17-20 (x =18.4), uppermost two and lowermost two or three raysunbranched, with about thirteen branched rays inbetween; pelvic fin rays I 5, no fraenum, full basalmembrane; fifth pelvic fin ray branched multiple times(variable), 60-90 % (x = 70 %) length of fourth ray. Lat-eral scales 23-25 (x = 23.7) (14.0 mm SL with 21), ante-rior transverse scales 7-10 (x = 8.3), posterior trans-verse scales 6-9 (x = 7.8), predorsal scale rows 8-12 (x = 10.6) (1 1.1 mm SL specimen with 4); scales onopercle, pectoral base, and prepelvic region cycloid; allother body scales ctenoid; 0-3 (x = 1.5) scales presenton anterodorsal corner of the opercle; 0-3 (x = 2.1)scale rows on the pectoral base; prepelvic region with4-7 (x = 5.2) scale rows; specialised scale on pelvicbase, with an expanded distal margin resembling a fan.Teeth in both upper and lower jaw with large, curved,spaced, canines in outer row , with smaller irregularinner rows of conical teeth, and innermost row of lowerjaw with spaced canines about half the size of outer;tongue rounded to bilobed. Gill opening to below mid-pupil; gill rakers on first arch 0 + 12-14 (n = 10, x = 0 +13.0). Anterior nasal opening tubular , slightly flared;posterior nasal opening a pore with raised rim, adnateto anterior margin of eye. A pair of thin fleshy lappets,half a pupil width apart, a quarter of a pupil widthbehind frontal ridge, on either side of dorsal midline(Fig. 10).

Trenches: trenches as in Trimma lantana . Trimmamendelssohni has a frontal ridge with an overall trian-gular shape (apex pointing anteriorly , with convex tip),in dorsal profile, similar to that of Trimma caesiura (seeFig. 1E). The apex of the frontal region may slopesteeply, anteroventrally into a narrow trench betweenthe orbits, while in other specimens (about 50%, 17.0mm SL and smaller) the apex slopes gradually (with no

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Fig. 8. Trimma mendelssohni, 17.9 mm SL, freshly-collected, female, Comores, ROM 59789. Photo by R. Winterbottom.

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ridge) into a wide space between the orbits. Where theapex slopes steeply between the orbits, the sides of thefrontal ridge end abruptly behind the orbits, forming theposterior wall of narrow, deep postorbital trenches. Theposterior wall of the postorbital trenches (normallyformed by the frontal ridge ending abruptly behind theorbits) is absent when the frontal apex slopes graduallybetween the orbits, thus forming wide, shallow postor-bital trenches.

Colour in life (based on underwater photos by G.Barrall, freshly-collected specimens, based on colourslides, from the Comores) : the body colour consists ofbrown/red background with five poorly-defined whitevertical bars, fading ventrally, extending from just ante-rior to the origin of the first dorsal to the mid peduncle.

The ground colour of the head is mottled brown/red,and is speckled with numerous melanophores andchromatophores. Three white bars dotted with chro-matophores occur under the orbit. The first, a quarterpupil width, is situated under the anterior half of theorbit and is separated from the second by a thick brown/ red blotch under the mid-orbit, representing theground colour. The second white bar, about a quarter toa half pupil width, is situated under the posterior half ofthe pupil, and is followed by a similar , irregular brown /red blotch. A third irregular white bar , nearly a pupilwidth, passes vertically across the vertical limb of thepreopercle (see Fig. 9). The other head blotches grad-ually become larger posteriorly , and become less dis-tinct along the nape and trunk. The scale pockets of the

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Fig. 10. Trimma mendelssohni, live, showing white nape lappets, Egyptian Red Sea. Photo by G. Barrall.

Fig. 9. Trimma mendelssohni. Underwater photograph of live specimen, Egyptian Red Sea. Photo by G. Barrall.

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nape and trunk have strong brown/red outlines, withlarge white centres (see Fig. 9). The first white bar onthe body is half a pupil in width and is found at the ori-gin of the first spine of the first dorsal fin (dorsal halfbright white in life), before fading ventrally behind thepectoral base. The second bar , approximately a halfpupil width, begins at the origin of the fifth dorsal spine(off-white in life), and usually fades ventrally behind thepectoral rays, rarely reaching the ventral margin (seeFig. 9). The third bar is situated on the dorsum at theorigin of the spine of the second dorsal fin (dorsal halfbright white in life), fades ventrally , and often reachesthe ventral margin. The fourth and fifth bars, both half apupil in width, are bright white in life, and reach the lat-eral mid-line. The fourth bar is located at the origin ofthe last ray of the second dorsal, while the fifth islocated on the dorsal half of the mid-peduncle (see Fig.9). The first dorsal, pectoral and pelvic fins are tingedred and hyaline distally. The second dorsal fin has rowsof red spots fading distally , with hyaline segments inbetween each row . The caudal fin has rows of faintbrown / red spots, which fade posteriorly along the hya-line rays. Trimma mendelssohni has a pair of fleshylappets on the dorsal surface of the head, which arebright white in life (see Fig. 10), but become dark brownwhen preserved in 70% ethyl alcohol.

Colour in alcohol: the body colour is a pale strawyellow with scale pockets margined with chro-matophores. The blotches and pale sections under theorbit reflect the facial pattern in life. The anterior half ismore intensely covered in melanophores than posteriorhalf. Along the dorsum, there are pale bars that endshort of the lateral mid-line.

Affinities and DiscussionTrimma mendelssohni possesses several features

unique to Trimma, allowing for easy identification bothin the lab and in the field. These features include a pairof fleshy lappets on the dorsal surface of the head thatare bright white in life (dark brown when preserved –

Fig. 1E); a specialised pelvic scale that has anexpanded distal end resembling a fan; an average of 11predorsal scale rows; a complex branching fifth pelvicfin ray (appearing bushy); a full basal membrane; abrown/red ground colour with white bars along thebody. These features are in agreement with Winterbot-tom’s 1995 brief description of T. mendelssohni withrespect to the fleshy lappets on the nape, the status ofthe trench and the absence of gill rakers on the first gillarch. These features help distinguish T. mendelssohnifrom the group consisting of T. lantana, T. naudei andT. caesiura, which have a single dichotomous branch inthe fifth pelvic ray; 18 or fewer pectoral rays; less than10 predorsal scale rows; a red / orange body colourwith head blotches and white body spots; and a poste-rior nostril separate from the orbit.

It was observed that the specimens with the mostelongate spines were males, but correlations betweensex, SL, and locality with first dorsal spine length werenot apparent. This could have been confounded bythe low number of males compared with females col-lected across sites.

DistributionTrimma mendelssohni is currently known from the

Comores, Egypt, and the Gulf of Aqaba, Mauritius,Madagascar, Rodriges, Seychelles, Sudan, andYemen. Specimens were collected at a depths rangingfrom 2-20 m, over coral flats rising above the rubble /sandy bottom.

Trimma winterbottomi Randall et al.(Figs. 1C, 11-13)

Trimma winterbottomi Randall et al., 1994: 250 (Per-sian Gulf) (replacement name)

Gobius townsendi Boulenger, 1897: 421 (Mekran Coast)– junior subjective homonym of Gobius townsendiEigenmann and Eigenmann 1888: 463 (San Diego).

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Fig. 11. Trimma winterbottomi, 24.2 mm SL, freshly-collected, female, Persian Gulf, BPBM 30462. Photo by J. E. Randall.

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orbits that slopes steeply into a wide interorbital trenchanteromedially and into postorbital trenches laterally(see description below). Posterior nasal openingadnate to eye; no elongate dorsal spines; 10 or moreelements in second dorsal; fifth pectoral ray with twosequential branches (occasionally one dichotomousbranch); 18 or more pectoral rays; lacking scales onhead; body colour orange / red with four white bars ondorsum, and light orange blotches on white colouredhead and nape; dark triangular group of melanophoresabove opercle.


17 females and 1 cleared and stained specimens, 10.8-24.1 mm SL. Vizhimjam, Kerala State, India, 1 (20.8),BPBM 27723; Sur, Oman, 1 (13.6), ROM 40020; 13 kmeast of Sur , Oman, 3 (10.8-1 1.1) ROM 40005; JanaIsland, Saudi Arabia, 1 (24.1), BPBM 30462; Galle, SriLanka, 2 (14.8-14.9) ANSP 152760; Hikkaduwa, SriLanka, 2 (16.8-19.2) ANSP152745; 2 (13.9-15.8)USNM 339544; Trincomalee, Sri Lanka, 2 (18.1-19.8)USNM 295227; Phuket, Thailand, 2 (19.1-20.4) ROM68103; 1 (18.8) ROM 68743; one cleared and stained,(20.4), ROM 1744 CS.

Dorsal fins VI + I 10-1 1(x = 10.4), no elongate dorsalspines; anal fin I 9-10 (x = 9.9); pectoral fin rays 18-22(x = 20.1), uppermost two unbranched, with about ninebranched rays in between (number of lowermostunbranched not determined); pelvic fin I 5, no fraenum,basal membrane normally complete, occasionally ves-tigial; fifth pelvic fin ray branched twice sequentially(occasionally once dichotomously), 50-95% (x = 73.8%) length of fourth ray. Lateral scales 24-29 (x = 26.8),anterior transverse scales 8-1 1 (x = 10.3), posteriortransverse scales 8-11 (x =10.4), no scales on predor-sal, opercle, pectoral base, or prepelvic region; bodyscales ctenoid. Teeth in both upper and lower jaw with

large, curved, spaced, canines in outer row, withsmaller irregular inner rows of conical teeth, and inner-most row of lower jaw with spaced canines about halfthe size of outer; tongue rounded to bi-lobed. Gill open-ing to below the mid pupil; gill rakers on first arch 1-3 +13-14 (n = 10, x = 2.3 + 13.8). Anterior nasal openingtubular, slightly flared; posterior nasal opening a porewith raised rim, adnate to anterior margin of eye.

Trenches: trenches as in Trimma lantana . Trimmawinterbottomi has a frontal ridge with an overall trian-gular shape (apex pointing anteriorly , with pointed tip),in dorsal profile (see Fig. 1E). In the frontal midline, theapex of the ridge slopes steeply , anteroventrally into adeep trench between the orbits. Behind the orbits, theridge abruptly ends forming the posterior wall of deep,narrow postorbital trenches. Smaller specimens (13.6mm SL and smaller) have an apex that slopes gradu-ally (with no ridge) into a wide space between the orbits

Colour in life: (freshly collected specimens, basedon colour slides from Thailand and the Persian Gulf):the body colour is orange/red with five poorly defined,white, elongated spots along the dorsum, between theposterodorsal margin of the peduncle and the origin ofthe first dorsal fin. The head and nape are white withelongated, light orange blotches. A single light orangebar, half a pupil width, occurs under the anterior half ofthe orbit, followed by an irregular light orange blotch,about pupil width, under the posterior half of the orbit,about half a pupil width apart from each other (seeFig. 11). The posterodorsal margin of the upper jawoccasionally has a thin, faint, vertical line ofmelanophores, about one third pupil length (seeFig. 12). A dark triangular group of melanophoresoccurs half a pupil width anterior to the posterodorsalmargin of the opercle (see Fig. 1 1, 12). The elongatedlight orange blotches on the head and nape becomeless distinct posteriorly , darken, and form an orange /red ground colour on the trunk. The scale pockets arenot outlined in melanophores and are masked by theground colour. The anterior off-white background

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Fig. 12. Trimma winterbottomi, 20.9 mm SL, freshly-collected, female, Phuket, Thailand, ROM 68091. Photo by R. Winterbottom.

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colour is reduced to irregular bars in between the coa-lesced orange blotches, and gradually decreasesbecoming smaller posteriorly. The most developed off-white stripe is an exaggerated zigzag in shape, andpasses along the dorsolateral surface of the trunk. Thefirst white spot, nearly pupil width, is situated on the ori-gin of the second dorsal fin, and is somewhat verticallyelongated, extending ventrally for a similar distance.The second white spot, also nearly pupil width, occursat the origin of the fourth ray of the second dorsal, andis somewhat elongated, at times extending 2 pupilwidths ventrally. The third white spot, similar to the sec-ond, occurs at the origin of the last ray of the seconddorsal fin. The fourth white spot, slightly larger thanpupil width, is situated on the dorsum of the mid-pedun-cle. A fifth, diffuse white bar , about half the size of thefourth, is situated on the posterodorsal corner of thepeduncle, and is rarely observed in freshly-collectedspecimens (see Fig. 11).

Colour in alcohol: the body colour is a pale strawyellow. The head and body pattern of freshly-collectedspecimens is retained in preserved specimens, whereblotches appear as groups of melanophores. The ven-tral midline of most female specimens has 0-2 (x = 1.1)melanophores on the prepelvic region, which are lack-ing in males.

Affinities and DiscussionTrimma winterbottomi and T. mendelssohni share fea-

tures not found in T. naudei, T. lantana or T. caesiura.Both T. winterbottomi and T. mendelssohni have fullbasal membranes (occasionally vestigial for T. winter-bottomi), usually more than one dichotomous branch inthe fifth pelvic ray, adnate posterior nasal openings and8 or more second dorsal rays (vs. a vestigial mem-brane, one dichotomous branch in the fifth pelvic, pos-terior nasal opening separate from the eye, and 8 orfewer second dorsal rays). Trimma winterbottomi is

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Fig. 13. Spot distribution map for the species of Trimma: T. caesiura – circles; T. lantana – triangles, T. mendelssohni –squares; T. naudei – diamonds; T. winterbottomi – inverted triangels.

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unique to this group in that it is the only speciesdescribed here that has no head scales, has 9 or morerays in the second dorsal and has an anal fin ray countof 9 or more. This species also possesses a triangulargroup of melanophores above the opercle, and a dis-tinct preserved banding pattern across the nape andtrunk, which is retained in preserved specimens.

The occurrence of prepelvic melanophores in pre-served specimens may be sex and / or location spe-cific. It was observed that females from Sri Lanka andOman possess the prepelvic melanophores (nofemales were collected from Kerala, India). Specimensof both sexes collected from Saudi Arabia and Thailandlack the prepelvic melanophores. More specimens,particularly males, would be required before any con-clusive analysis can be performed to support or refutesexual dimorphism. However, the total collected mate-rial of 3 males to 17 females may indicate that thisspecies forms harems, as has been reported forT. okinawae by Sunobe and Nakazono (1990) andSunobe and Nakazono (1993).

DistributionTrimma winterbottomi is currently known from India,

Oman, the Persian Gulf, Saudi Arabia, Sri Lanka, andwestern Thailand. Specimens were collected at depthsranging from 5-15 m, over coral and within the reef /rubble slope.

AcknowledgementsMany thanks to all the curators and collection man-

agers who have assisted R. W’ s research of Trimmaover the last two decades. G. R. Allen, G. Barrall, J. E.Randall and H. Nagano are thanked for permission touse their photographs. Rick Winterbottom gratefullyacknowledges the financial contributions of the ROMFoundation, the Centre for Biodiversity and Conserva-tion Biology, and a National Sciences and EngineeringResearch Council of Canada research grant (OGP0007619). The study published herein represents Con-tribution No. 263 of the Centre for Biodiversity and Con-servation Biology of the Royal Ontario Museum to thebiological sciences.

ReferencesAkihito, Prince Hayashi, M., & T. Yoshino. 1988.

Suborder Gobioidei. In: The Fishes of the JapaneseArchipelago second edition. (Eds. H. Masuda, K.Amaokia, C. Araga, T. Uyeno & T. Yoshino): 228-289, 445; pls. 235-258, 375. Tokai University Press,Tokyo.

Allen, G. R. 1997. Marine Fishes of T ropical Australiaand South-East Asia . W estern Australian Museum,Perth, Australia. 292 pp.

Boulenger, G. A. 1897. Descriptions of new fishesfrom the Mekran Coast, Persia. Annals and Magazineof Natural History, 6: 420-422.

Burgess, W . E., Axelrod, H. R. & R. E. Hunziker .

1990. Dr. Burgess’s Atlas of Marine Aquarium Fishes(Second Edition). T.F.H. publications, New Jersey.768 pp.

Goren, M. 1978. A new gobiid genus and seven newspecies from Sinai coasts (Pisces: Gobiidae).Senckenberg Biologica, 59 (3/4): 191-203.

Hubbs, C. L. & K. F. Lagler. 1958. Fishes of the GreatLakes Region . University of Michigan Press, AnnArbor, Michigan. 213 pp.

Jordan, D. S. & A. Seale. 1906. The fishes of Samoa.Description of the species found in the archipelago,with a provisional check-list of the fishes of Oceania.Bulletin of the U.S. Bureau of Fisheries, 25: 173-455.

Kuiter, R. H. 1992. Tropical Reef-Fishes of theWestern Pacific – Indonesia and Adjacent W aters.Penerbit PT, Jakarta, Indonesia. 314 pp.

Randall, J. E . 1998. Zoogeography of shore fishes ofthe Indo-Pacific region. Zoological Studies , 37 (4):227-268.

Randall, J. E., Downing, N., McCarthy , L. J., Stana-land, B. E. & A. B. Tarr. 1994. Fifty-one new recordsof fishes from the Arabian Gulf. Fauna Saudi Arabia,15: 220-258.

Smith, J. L. B. 1957. The fishes of Aldabra (Part VI).Annals and Magazine of Natural History, 12: 817-829(for 1956).

Sunobe, T. & A. Nakazono. 1990. Polygynous matingssystem of Trimma okinawae (Pisces: Gobiidae) atKagoshima, Japan with a note on sex change.Ethology, 84: 133-143.

Sunobe, T. & A. Nakazono. 1993. Sex change in bothdirections by alteration of social dominance in Trimmaokinawae (Pisces: Gobiidae). Ethology, 94: 339-345.

Winterbottom, R. 1984. A review of the gobiid fishgenus Trimma from the Chagos Archipelago, centralIndian Ocean, with the description of seven newspecies. Canadian Journal of Zoology, 62: 695-715.

Winterbottom, R. 1995. Red Sea gobiid fishes of thegenus Trimma, with the description of two newspecies. Revue française d’Aquariologie , 22 (3-4):93-98.

Winterbottom, R. & M. Burridge. 1992. Revision ofEgglestonichthys and of Priolepis species possessinga transverse pattern of cheek papillae (T eleostei;Gobiidae), with a discussion of relationships. Cana-dian Journal of Zoology, 70: 1934-1946.

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KeywordsStomach contents, Synodus saurus , seasonal

variations, growth changes, feeding strategy , preyorientation

AbstractThe feeding habits of the lizardfish Synodus saurus,

were studied in the Azores archipelago, north-easternAtlantic. Factors examined were diet composition,prey importance, season, fish size, feeding strategyand prey orientation in the oesophagus. The stomachcontents of 308 specimens were collected and ana-lyzed between March and November 2000. Synodussaurus is common in Azorean waters. Though itprefers small gregarious pelagic fish, it also feeds onepibenthic and benthic prey . W e found the followingprey in its diet: 9 families of teleostean fishes (Carangi-dae, Clupeidae, Cynoglossidae, Gobiidae, Labridae,Myctophidae, Sparidae, Sphyraenidae and Synodonti-dae), two families of crustaceans (Cymothoidae andScyllaridae) and one of cephalopods (Loliginidae). TheEuropean pilchard, Sardina pilchardus was the com-monest prey. The diet of the lizardfish varies with theseason, showing the greatest diversity (given by theShannon-Wiener index) in October . There is a signifi-cant correlation with the sea temperature. The size ofthe predator is not correlated to the size of its prey .However, we observed a positive correlation betweenthe size of the predator and the quantity of food in itsstomach. The orientation of the prey in the oesopha-gus may partly depend on the predator ’s size. How-ever, analysis of the stomach contents provided noinformation on the selection of prey.

ZusammenfassungDas Fressverhalten vom Eidechsenfisch Synodus

saurus wurde im Azoren Archipel (nordöstlicherAtlantik) erforscht. Die untersuchten Faktoren waren:Zusammensetzung der Nahrung, Wichtigkeit derBeutetiere, Jahreszeit, Fischgröße, Fressstrategie undOrientierung der Beute in der Speiseröhre. In der Zeitvon März bis November 2000 wurde der Mageninhaltvon 308 Exemplaren gesammelt und analysiert. Syn-

odus saurus ist in den Gewässern der Azoren weit ver-breitet. Obwohl diese Art kleine pelagische Schwarm-fische vorzieht, frisst sie auch auf Substraten desGewässerbodens lebende- sowie direkt auf demBoden lebende Beutetiere. Wir fanden die folgendeBeutetierarten in ihrer Nahrung: 9 Familien von Teleost-fischen (Carangidae, Cynoglossidae, Clupeidae,Gobiidae, Labridae, Myctophidae, Sparidae,Sphyraenidae und Synodontidae), zwei Familien vonKrebstieren (Cymothoidae und Scyllaridae), sowie eineCephalopoden-Familie (Loliginidae). Die EuropäischeSardine, Sardina pilchardus war das am meisten vor-kommende Beutetier . Die Nahrung der Eidechsen-fische variiert jahreszeitlich und zeigt die unter-schiedlichste Mannigfaltigkeit (nach dem Shannon-Wiener Index ) im Oktober . Es besteht eine bedeu-tende W echselbeziehung mit der Meerestemperatur .Die Größe des Räubers korreliert nicht mit der Größeseiner Beute. Wir haben jedoch eine positive Korrela-tion zwischen der Größe des Räubers und derNahrungsmenge in seinem Magen feststellen können.Die Orientierung der Beute in der Speiseröhre hängtteilweise mit der Größe des Räubers zusammen.Jedoch Analysen des Mageninhalts ergaben keineInformationen über die Auswahl der Beute.

RésuméLes habitudes alimentaires du poisson-lézard

Synodus saurus ont été étudiées dans l'archipel desAçores, au nord-est de l'Atlantique. Facteurs analysés:composition du régime, importance des proies, saison,taille des poissons, stratégie alimentaire et orientationde la proie dans l'oesophage. Le contenu stomacal de308 spécimens a été collecté et analysé de mars ànovembre 2000. Synodus saurus est commun dans leseaux des Açores. Malgré sa préférence pour de petitspoissons pélagiques grégaires, il se nourrit aussi deproies épibenthiques et benthiques. Nous avons iden-tifié les proies suivantes dans le régime: 9 familles depoissons téléostéens (Carangidae, Clupeidae, Cyno-glossidae, Gobiidae, Labridae, Myctophidae, Sparidae,Sphyraenidae et Synodontidae), deux familles de crus-tacés (Cymothoidae et Scyllaridae) et une de cépha-

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Feeding habits of the lizardfish Synodus saurus (Linnaeus, 1758)(Actinopterygii: Synodontidae) from the Azores

Marta S. C. Soares, Luis Sousa and João Pedro Barreiros1

1) Departamento de Ciências Agrárias1, Universidade dos Açores. PT-9701-851 Angra do Heroísmo, Portugal E-mail: [email protected] (corresponding author)


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lopodes (Loliginidae). Le pilchard européen, Sardiniapilchardus, était la proie la plus fréquente. Le régime dupoisson-lézard varie en fonction de la saison, avec unpic de diversité (selon l'index Shannon-Wiener) enoctobre. Il existe une corrélation significative avec latempérature de la mer. La taille du prédateur n'est pascorrélée à celle de sa proie. Pourtant, nous avons con-staté une corrélation positive entre la taille du prédateuret la quantité de nourriture dans son estomac. L'orien-tation de la proie dans l'oesophage peut dépendre enpartie de la taille du prédateur . Néanmoins, l'analysedes contenus stomacaux ne donne pas d'informationsur la sélection des proies.

SommarioSono descritte le abitudini alimentari del pesce

lucertola Synodus saurus nell’arcipelago delleAzzorre, Atlantico nord-orientale. I fattori esaminatisono stati: la composizione della dieta, l’importanzadella preda, la stagione, le dimensioni, la strategia el’orientamento della preda nell’esofago. È stato anal-izzato il contenuto gastrico di 308 esemplari raccoltinel periodo marzo-novembre 2000. Synodus saurus èpiuttosto comune nelle Azzorre. Sebbene preferiscapiccoli pesci pelagici gregari, si nutre anche di speciebentoniche e epibentoniche. Sono state individuate leseguenti prede: 9 famiglie si pesci teleostei (Carangi-dae, Clupeidae, Cynoglossidae, Gobiidae, Labridae,Myctophidae, Sparidae, Sphyraenidae e Synodonti-dae), due famiglie di crostacei (Cymothoidae e Scyl-laridae) e una di cefalopodi (Loliginidae). La sardinaeuropea, Sardina pilchardus, era la preda piùcomune. La dieta di S. aureus cambiava con le sta-gioni, con una maggiore diversità (indice di Shannon-Wiener) in ottobre. Si osservava inoltre una corre-lazione significativa con la temperatura del mare. Lataglia del predatore non risultava invece correlata conquella della preda. Tuttavia, si osservava una corre-lazione positiva tra la dimensione del predatore e laquantità di cibo nello stomaco. L ’orientamento dellapreda nell’esofago può in parte dipendere dalla tagliadel predatore. Tuttavia, l’analisi del contenuto dellostomaco non ha fornito informazioni sulla selezionedella preda.

IntroductionThe lizardfish, Synodus saurus (Linnaeus, 1758) is a

common epibenthic fish in the waters around theAzores (Santos et al. , 1997). Its feeding habits haveonly been studied in the Mediterranean (Golani,1993). More recently , studies on its predatory andagonistic behaviour (Soares et al. , 2002) and itsreproduction (Sousa et al. , 2003) have been carriedout on populations in the Azores. The species occursin Madeira, the Canaries, Cape V erde, the Mediter-ranean, and of f the Moroccan coast. In the westernAtlantic it is known from Bermuda, the Bahamas andthe Leeward Islands (Sulak, 1986).

Synodus saurus is commonly found on sandy bot-toms in depths from less than 20 m down to 400 m(Sulak, 1983). It has a small head with medium-sizedeyes, and its colour pattern (Fig. 1) provides a veryeffective camouflage. Despite spending most of itstime buried in the sand, the lizardfish is a highlymobile predator, capable of capturing pelagic fishes inmidwater (Soares et al. , 2002). In the Azores, itsreproduction is continuous, peaking in the summer(Sousa et al., 2003).

This work describes the diet of Synodus saurus inthe Azores and discusses data on diet shifts, the rela-tionship between predator size and food taken, andthe orientation of prey in the oesophagus. It is one ofa series of papers describing the diet of severalcoastal species from the Azores, aimed at achievingbetter understanding of food webs and predator-preyrelationships (e.g., Barreiros & Santos, 1998; Moratoet al., 2000; Barreiros et al., 2002, 2003).

Materials and MethodsStudy area and sampling procedure: S. saurus

was sampled around Terceira Island, in the Azores(Fig. 2). A total of 308 fish were collected by spearfishing between March and November, 2000. This col-lection method has the advantage of avoiding theregurgitation of stomach contents (Bowen, 1983;Morato et al. , 2000), and facilitating the selection ofspecimens within a given species since it avoids anyby-catch (Derbal & Kara, 1996).

The specimens were frozen and measured to thenearest millimetre (total length-TL) and weighed to thenearest 0.1 g. The digestive tract was removed andthe fullness of the stomach recorded using a semi-quantitative scale. The stomachs were fixed in 10%buffered formalin and preserved in 70% alcohol. Theempty stomachs and stomach contents wereweighed. Food items were identified down to the low-est possible taxonomic level. To minimize the under-estimation of small, soft prey , the utmost care wasgiven to the identification of even the smallest frag-ments. Prey items were weighed to the nearest 0.01g after removing the surface water and, when undi-gested, items were measured (L) and allocated to alength group. The number of items of prey and theirorientation in the oesophagus were also recorded.Stomach contents that contained more than one typeof prey were described as “mixed” if the orientation ofthe prey differed (L’Abée-Lund, 1996). The orientationof prey in advanced states of digestion was impossi-ble to establish and was described as “undetermined”.

Stomach content analysis: The quantitative impor-tance of dif ferent prey in the diet was expressed asfollows (Berg, 1979; Hyslop, 1980):

Vacuity index (VI), as the percentage of empty stom-achs;

Repletion index (RI), as the percentage of stomachswith food;

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Feeding habits of the lizardfish Synodus saurus (Actinopterygii: Synodontidae) from the Azores

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Marta S. C. Soares, Luis Sousa and João Pedro Barreiros

Fig. 1. Camouflaged specimen of S. saurus. Photo by Peter Wirtz, ©imagDOP

Table I. Percentage contribution of prey groups and species to the diet of S. saurus.

Prey category n Cn Cw F IRI

FishSardina pilchardus 196 32.9 31.6 42.8 2759.7Sphyraena viridensis 67 11.3 1.2 4 49.5Pagellus acarne 9 1.5 6.3 3.5 27.3Coris julis 11 1.8 12.7 4 58Symphodus mediterraneus 1 0.2 3.4 0.5 1.8Trachurus picturatus 3 0.5 7.1 1.5 11.3Gobius paganellus 4 0.7 1.1 2 3.6Synodus saurus 3 0.5 7.1 1.5 11.4Symphurus nigrescens 4 0.7 0.1 1 0.8Mullus surmuletus 1 0.2 0.1 0.5 0.1Unidentified Myctophidae 2 0.3 0.1 0.5 0.2Unidentified Labridae 2 0.3 0.3 1 0.7Unidentified Sparidae 1 1.7 2.2 0.5 1.2Unidentified Pleuronectiformes 1 0.2 0.05 0.5 0.1Unidentified teleostei 260 43.6 11.6 11.6 3405.8

Total fish 565 96.3 84.9 125.5 6331.5

CrustaceaAnilocra physodes 28 4.7 0.6 2.8 45.5Scyllarus arctus 1 0.2 0.06 0.01 0.1

Total crustaceans 29 4.9 0.7 2.8 45.6

CephalopodaLoligo forbesi 2 0.3 0.7 1 1

Total cephalopods 2 0.3 0.7 1 1

No. of stomachs examined 308No. of empty stomachs 107Mean fish total length (mm) 281.5Mean stomach content weight (g) 3.9Mean number of prey items per stomach 1.9

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Percent frequency of occurrence ( F), based on thenumber of stomachs in which a food item was found,expressed as a percentage of the total number ofnon-empty stomachs;

Percent numerical abundance ( Cn), the number ofeach prey item in all non-empty stomachs in a sam-ple, expressed as the percentage of the total numberof food items in all stomachs in a sample;

Percent gravimetric composition ( Cw), the wetweight of each prey item, expressed as the percent-age of the total weight of the stomach contents in asample.

Importance of prey items was identified using theindex of relative importance ( IRI) of Pinkas et al.(1971): IRI = (Cn + Cw) × F.

The IRI is known to bias results in some cases(Tirasin & Jorgensen, 1999). However, since one preywas clearly dominant in the sample, it was justifiablein our case.

Niche breadth for the utilization of food resourceswas calculated according to the Shannon-Wienerindex (Krebs, 1989; Labropoulou et al., 1998):

H´ = ∑ (pi log epi)

where pi is the proportion of a specific prey categoryfor the n categories of prey listed. The Shannon-Wiener index value increases with the number ofspecies. S pecies e venness w as d etermined w ithPielou´s evenness index (Pielou, 1977):

J`= H´/loge(S)where H´ is the Shannon-Wiener index and S is the

total number of species. Correlations (Spearman rank) were tested between

different pairs of variables: vacuity index and water

temperature, mean stomach content weight andpredator length, mean prey length and predator lengthand mean prey weight and predator length. All theprevious variable relationships were also tested bysimple linear regression (Zar , 1996). Dif ferences inprey orientation in the oesophagus were tested withthe Sign-Test, according to the non-parametric char-acteristics of the studied data (Ludwig & Reynolds,1988; Krebs, 1989).

Seasonal changes in diet were analyzed by the ninesampling months: March (n = 14), April (n = 21), May(n = 42), June (n = 36), July (n = 23), August (n = 41),September (n = 56), October (n = 47) and November (n = 28). The total lengths of the fish examined rangedfrom 150 to 460 mm; the mean length was 281.5 andthe standard deviation (SD) 55.4. V ariations withinfish size were examined by allocating predatorlengths to five size groups (TL): 150-199 mm (n = 16);200-249 mm (n = 85); 250-299 mm (n = 84); 300-349mm (n = 82); > 350 mm (n = 41).

Feeding strategy was calculated using the newAmundsen et al. (1996) approach to the Costello(1990) method, which combines the frequency ofoccurrence (Fi) of a given prey (expressed by thenumber of stomachs in which that prey occurs) andthe prey specific abundance (Pi) (as the percentage ofa prey in only those predators in which it occurs):

Fi = (Ni /Nt) × 100Pi = (ΣSi /ΣSt) × 100

where Ni is the number of predators with prey i in theirstomach, N is the total number of predators with stom-ach contents, Si the number of stomachs with prey i,and St the total prey in the stomachs containing preyi. The percent abundance, increasing along the diag-onal from the lower left to the upper right corner , pro-

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Fig. 2. The Azores archipelago, north-east Atlantic.

S

i = l

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temperature, for the analyzed months (Fig. 4) areweakly correlated (rs = 0.74; p < 0.05).

The median number of prey found in each stomachwas higher in the first months of sampling. Thespecies total was greatest in October . The monthlyweight was quite variable but also reached a maxi-mum in October (Table II).

Diet diversity was higher in the later samplingmonths, particularly October (H´= 0.758) and August(H´= 0.675). The last sampling month showed lowerdiversity ( H´= 0.326). The evenness distribution was similar to H’. Its maximum value was observed in October (J´= 0.785), and its minimum in Novemb-er (J´= 0.466). Neither diversity nor evenness values were correlated with the water temperature (p > 0.05).

Diet vs. fish size: Although not significantly corre-lated (p > 0.05), the repletion index (RI) shows someontogenetic variation. In the first two length groups(100-149 mm and 150-199 mm) it was almost thesame, and the highest value was reached in the 250-299 mm group.

The mean weight of the stomach contents increasedwith the size of the predator, up to a length of 330 mm,when it reached a maximum (Fig. 5). Mean stomachcontent weight increased as the fish grew (predatorTL) with a significant positive linear relation (r2 = 0.91;rs = 0.82; p < 0.05). Also, the frequency with whicheach prey species was found in the stomachs differedwith predator length (Fig. 6). Maximum prey fre-quency was recorded in the 249 – 299 mm group, with275 items. Sardina pilchardus was found in most ofthe groups.

Predator-prey length and weight relationship:The minimum prey size was constant whatever the


Fig. 3. Specific prey frequency per month.

vides a measure of prey importance, with dominantprey at the upper and end, rare or unimportant prey atthe lower end. The vertical axis represents the feed-ing strategy of the predator, in terms of specializationor generalization (Amundsen et al., 1996).

Results Diet composition and prey importance: Of the

total 308 stomachs analyzed, 201 (65.3 %) containedfood. In these, 596 prey items belonging to threegroups (fish, crustaceans and cephalopods) wereidentified. Fish (13 species) were the most importantgroup.

The European pilchard, Sardina pilchardus was themost important p rey, w hile t he y ellowmouth b ar-racuda, Sphyraena viridensis, and the parasitic iso-pod, Anilocra physodes, were also found in high num-bers (Table I). However, their weight importance wascomparatively low. The labrid Coris julis (L.) was thesecond most important prey according to its Cw value.The species A. physodes , besides being part of thispredator’s diet, is also an ectoparasite and individualsranging in size from 2 to 35 mm were found attachedto S. saurus’ mouth at a frequency of 1.3 (SD = 0.06)per fish. The relative importance of the prey groupsand prey species according to the calculated valuesof the IRI are also given in Table I.

Seasonal variations in prey composition: Thedistribution of food items varied with the season (Fig.3). Initially, the frequency increased, with a maximumvalue in April (133 items). Only one species, S.pilchardus, was found in all samples. Sphyraena viri-densis occurred only in the first two months. Diversityincreased in the final months.

The values of the vacuity index (VI) and sea water

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size of the predator (Fig. 7). Maximum prey length didnot correspond to maximum predator size. Neverthe-less, a significant progression between these vari-ables can be seen.

Mean prey length increased with predator size (preda-tor SL), with a positive linear relation between both vari-ables (r 2 = 0.85; rs = 0.83; p < 0.05). As in the previouscorrelation, mean prey weight was also directly relatedto predator length. In this particular case, the linear rela-tion is highly significant ( r 2 = 0.90; rs = 0.94; p < 0.01).

Feeding Strategy: The species shown at upper leftin the diagram are consumed by a limited fraction of

the predators (Fig. 8), which indicates a highbetween-phenotype component. These are: C. julis ,Pagellus acarne , S. saurus , Trachurus picturatus ,Loligo forbesi , Symphodus mediterraneus , Symphu-rus nigrescens, unidentified Sparidae and unidentifiedPleuronectiformes. The prey species shown lower leftin the diagram were occasionally eaten. Sardinapilchardus is the only dominant item in the lizardfish’sdiet (population specialization).

Prey orientation: S. saurus feeds on several typesof prey , with and/or without spines, pelagic and/orbenthic species. In all, 51% of S. saurus´s prey wereeaten tail-first. Only 13.7% were eaten head-first.However, some species were always eaten head-first:S. mediterraneus, Gobius paganellus, S. saurus andL. forbesi (Table III). The majority of stomachs con-tained one (27.6%), two (8%) or three (5.6%) prey, butup to 30 items were recorded.

DiscussionThis study indicates that the lizardfish from the

Azores archipelago feeds primarily on small gregari-ous pelagic fish, such as S. pilchardus, S. viridensisand T. picturatus. The presence of juvenile S. viriden-sis in the stomach contents of an epibenthic specieshas never been reported before. S. saurus also feedson epibenthic and benthic species such as C. julis andP. acarne. Cannibalism, observed in this predator, is asurprisingly common form of predation in fish, pre-sumably because typically the young are so small


Table II. Seasonal variations in terms of mean number ofprey, mean weight (g) and the number of speciesrecorded in each month.

Table III. S. saurus: prey orientation in oesophagus.

Months Mean nº prey Mean prey weight Species per stomach per stomach (g) recorded

March 5 0.34 4April 6.34 0.27 4May 2.36 1.66 6June 2.55 0.96 5July 1.22 2.98 6August 0.97 0.99 9Septemb. 0.95 3.98 7October 0.74 5.53 10November 1.69 3.64 6

Orientation

Tail Head Undeterminedfirst first

Sardina pilchardus 139 32 24Sphyraena viridensis 39 8 20Pagellus acarne 8 -- --Coris julis 7 2 --Symphodus mediterraneus -- 1 --Trachurus picturatus 2 1 --Gobius paganellus -- 4 --Synodus saurus -- 3 --Symphurus nigrescens 1 3 --Mullus surmuletus 1 -- --Unidentified Myctophydae 2 -- --Unidentified Labridae 2 -- --Unidentified Sparidae 1 -- --Unidentified Pleuronectiformes 1 -- --Unidentified Teleostei 90 25 144Anilocra physodes 3 2 22Scyllarus arctus -- -- 1Loligo forbesi -- 2 --

Total 296 83 211

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compared with the parents (Moyle & Cech, 2000).Most of the time, predated individuals are very youngand are thus eliminated as potential competitors(Keenleyside, 1979). Juvenile S. saurus occupy thesame habitat as adults, thus becoming an easy prey .The capture of crustaceans and cephalopods by indi-viduals of this species or other Synodontidae hasbeen mentioned in some studies (Golani, 1993, Kagi-wara & Abilhôa, 2000). Budnichenko (1974) regis-tered the frequent occurrence of Loligo spp. in the

stomach contents of the confamilial species Sauridatumbil and S. undosquamis . The presence of anectoparasite (A. physodes) in the stomach contents ofS. saurus has never been reported before and will bediscussed elsewhere (Soares et al., in prep.).

Synodus saurus shows seasonal dif ferences in itsdiet composition. The relatively low values of theShannon-Wiener index throughout the samplingmonths, indicate that this predator’s diet is composedof few prey species. Only one species ( S. pilchardus)


Fig. 4. Vacuity index and sea water temperature, per month.

Fig. 5. Stomach contents mean weight (g) per length group.

Sto

mac

h co

nten

ts m

ean

wei

ght (

g)

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seems to have a positive association in its monthlydiet. The frequency of prey in the stomach alsochanges monthly. These data reflect environmentaldynamics and the abundance and distribution of thetarget prey species in a seasonal perspective.

The time required to digest the food in the stomachdecreases as temperature rises, due to increasedenzymatic activity (Jones, 1978). At higher tempera-tures, similar stomach contents are digested at higherrates (Knutsen & Salvanes, 1999). It is natural, there-fore, that the vacuity index is positively correlated to

the water temperature. This justifies the fact that onlya small fraction of the stomachs examined were com-pletely full and that most of the prey items werealready considerably digested.

Although changes to the predator, as it grows, justifypossible diet alterations, in the case of S. saurus thisis not proportional. The width and depth of the openmouth are linearly related to fish sizes and increasedbody and mouth sizes allow fish to capture a broaderrange of prey types and sizes (Labropoulou et al. ,1998). Also, as metabolic activity decreases with age


Fig. 6. Synodus saurus: distribution of prey frequency per size group.

Fig. 7. Relation between lengths of captured prey (mm) and predator length (mm).

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or size, it becomes more beneficial for larger fish toobtain more bulk by expending less energy(Labroupoulou et al. , 1998). Consequently , a largerpredator would have more food in its stomach. This isexactly the case with S. saurus. This positive relationenables the predator to capture a greater variety ofprey, with several competitive benefits. The relationbetween the length of predators and the length of itsprey may indicate that S. saurus exhibits selectivepredatory behaviour.

In terms of prey specific frequency , we noticed thatthe predator ’s middle size groups had higher abun-dance values. Nevertheless, the stomach contentsanalyses did not indicate a tendency towards thestraight selection of prey, in relation to the predator’slength, since all size classes appear to explore similarprey species. There is no distinct segregation in termsof prey items between dif ferent length groups.

S. saurus has a mixed feeding strategy , as it com-bines a population specialization towards S. pilcharduswith a high between-phenotype component. Mostpredators utilize several food sources simultaneously(individual specialization). Each species of this set ofprey is consumed by a limited fraction of the predators(Amundsen et al. , 1996). Milinski (1982) states that,when it comes to predation, predators with a minor suc-cess rate may not include preferential prey in their diet.These become generalists.

The hunting behaviour dif fers with prey species.According to Golani (1993), the prey orientation in thestomach may indicate the hunting strategy of thepredator (i.e. prey found head-first were most likely tohave been ambushed while those found tail-first wereprobably chased or caught from behind). Fast swim-mers, such as S. pilchardus and S. viridensis were cap-tured mainly by chasing, while S. saurus, G. paganel-

lus and S. mediterraneus were ambushed. Prey organ-isms were found pointing in both directions, suggestingthat ambush and chasing may be used for most of theprey found in S. saurus stomachs. L’Abée-Lund et al.(1996) affirm that the obvious advantage of head-firstorientation is that swallowing is easier and quicker andthat the advantage increases with increasing preylength because the prey is totally consumed. Whenswallowing a prey head-first, problems due to defen-sive mechanisms (e.g., spines, poison glands) are min-imized. This is because once the dorsal, anal and pec-toral spines are locked into place the prey can only betaken by a predator that can move its mouth around thespines (Moyle & Cech, 2000). The largest prey found inS. saurus stomachs was a 225 mm TL C. julis that hadbeen eaten head-first, agreeing with L’Abée-Lund et al.(1996). The observed data indicate that the orientationof the prey depends partly on its size. In this case, itsspeed and escape strategy would be determinant.Also, hunting strategy may change with predator size.However, analysis of the stomach contents did not indi-cate a clear trend in prey type selection according topredator size. Nevertheless, it is clear that an increasein size may favour an increase in predatory ability .

AcknowledgementsWe are grateful to Pedro Ré and Henrique Cabral of

the University of Lisbon. This work was made possi-ble by the support of the University of the Azores(Department of Agricultural Sciences) and the Univer-sity of Lisbon (Faculty of Sciences). Special thanksare due to Alfredo Borba, Carlos Vouzela, Goretti Bet-tencourt and Oldemiro Rego, for the use of laboratoryfacilities and support. Thanks are also due to the twoanonymous referees who helped to significantlyimprove this paper.


Fig. 8. Synodus saurus feeding strategy according to Amundsen et al. (1996).

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KeywordsKillifishes, Rio São Francisco, systematics, phy-

logeny, Rivulus, Rivulidae

AbstractRivulus paracatuensis n. sp. is described from a

small stream in the Rio Paracatú floodplains, Rio SãoFrancisco basin, Minas Gerais, Brazil. It was previouslymisidentified as R. decoratus, the only other species ofthe genus occurring in the São Francisco basin, butdescribed from Ibiraba, Bahia, which is about 1,000 kmnorth of the Rio Paracatú. The new species is easilydistinguished from R. decoratus by having more dorsal,caudal and pectoral fin rays, more vertebrae, morescales in the longitudinal series and more scale rowsaround the caudal peduncle, a wider basihyal, and adistinct male colour pattern.

ResumoRivulus paracatuensis n. sp., coletada em um

pequeno riacho na várzea do rio Paracatú, bacia do rioSão Francisco, Minas Gerais, Brazil, é descrita. Ela foiantes equivocadamente identificada como R. decora-tus, a única outra espécie do gênero que ocorre nabacia do São Francisco, mas descrita de Ibiraba,Bahia, que está cerca de 1.000 km a Norte do rio Para-catú. A nova espécie se distingue facilmente de R. dec-oratus por possuir mais raios nas nadadeiras dorsal,caudal e peitoral, mais escamas na série longitudinal emais séries de escamas em torno do pedúnculo cau-dal, basi-hial mais largo e um distinto padrão de col-orido em macho.

ZusammenfassungRivulus paracatuensis n. sp. wird von einem kleinen

Fluss im Überschwemmungsgebiet des Rio Paracatú,Rio São Francisco-Becken (Minas Gerais, Brasilien)beschrieben. Diese Art war vormals fälschlich als R.decoratus identifiziert, die einzige andere Art in dieserGattung, die zwar im Rio São Francisco-Beckenvorkommt, aber von Ibiraba, Bahia (ungefähr 1.000 kmnördlich vom Rio Paracatú), beschrieben wurde. Dieneue Art ist leicht von R. decoratur zu unterscheiden;sie hat mehr Rücken-, Schwanz- und Brustflossen-

strahlen, mehr Rückenwirbel, sowie mehr Schuppen inder Längsserie und auch um den Schwanzstiel herum.Ebenfalls anwesend ist ein breiteres Zungenbein,sowie ein anderes Farbmuster in männlichen Tieren.

RésuméRivulus paracatuensis n. sp. est décrit venant d'un

petit cours d'eau dans les zones inondables du RioParacatú, bassin du Rio São Francisco, Minas Gerais,Brésil. Il fut autrefois erronément identifié sous le nomde R. decoratus, la seule autre espèce du genre vivantdans le bassin du São Francisco, mais décrit d'Ibiraba,Bahia, qui se trouve à 1000 km au nord du Rio Para-catú. L'espèce nouvelle se distingue facilement de R.decoratus par un nombre plus élevé de rayons à la dor-sale, la caudale et aux pectorales, plus de vertèbres,plus d'écailles dans les séries longitudinales et plus derangées d'écailles autour du pédoncule caudal, unbasihyal plus large et un patron de coloration propreaux mâles.

SommarioRivulus paracatuensis n. sp. viene descritto da esem-

plari raccolti in un piccolo corso d’acqua dell’alveo delRio Paracatú, bacino del Rio São Francisco, MinasGerais, Brasile. Precedentemente confuso con R. dec-oratus, l’unica altra specie del genere presente nelbacino del Rio São Francisco, ma descritta ad Ibiraba,Bahia, a circa 1000 km a nord del Rio Paracatú. Lanuova specie si distingue da R. decoratus per avere unmaggior numero di raggi nelle pinne dorsale, caudale epettorali, più vertebre, più scaglie in linea longitudinale,più file di scaglie attorno al peduncolo caudale, ilbasiale più largo e una diversa livrea nel meschio.

IntroductionThe Rio São Francisco basin is the fourth largest river

basin in South America, its main course extending forabout 3,100 km. It drains an area of 631,133 km 2 innorth-eastern Brazil, mostly within the Caatinga, asemi-arid region. A high degree of fish diversity andspecies endemism has been recorded (e. g. Travas-sos, 1960; Britski et al., 1988), but until 1988 no rivulidhad been found in this basin. Surprisingly , recent


Rivulus paracatuensis n. sp. (Cyprinodontiformes: Rivulidae): a new rivuline species from the Rio São Francisco basin, Brazil

Wilson J. E. M. Costa

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


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Fig. 1. Geographic distribution of Rivulus paracatuensis and R. decoratus.

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research directed at sampling special habitats hasrevealed a diversified assemblage of endemic rivulids,with a total of 27 species (Costa, 2001, in press; seealso many other references in these two papers). How-ever, although the annual fish genera SimpsonichthysCarvalho and Cynolebias Steindachner are repre-sented by several species, some of which are wide-spread over a vast area and occur in many localities,Rivulus Poey is represented by a single species, R.decoratus Costa. This species is recorded from only afew localities in the São Francisco basin (Costa,1995a), contrasting with other species of Rivulusendemic to neighbouring river basins (i. e. Tocantinsand Paraná), where they are found at numerous sitesand in almost all shallow aquatic freshwater habitats.

Rivulus decoratus was first described on the basis ofspecimens collected in Ibiraba, northern Bahia (Costa,1989). Subsequently, R. decoratus was considered tobe a member of a clade termed the R. punctatus com-plex and was included in the taxonomic revision of thisgroup (Costa, 1995a). During the final stages of thepreparation of this paper , a new population was foundin the Rio Paracatú floodplains, western Minas Gerais,about 1,000 km in a straight line from the type locality(Fig. 1). Specimens of this population were at that timeidentified as R. decoratus (Costa, 1995a), despite hav-ing a distinct colour pattern. Subsequent morphologicalcomparison has provided strong evidence supportingrecognition of the population from the Paracatú flood-plains as a new species, described here.

Materials and methodsMeasurements and counts follow Costa (1995b).

Measurements are presented as percentages of stan-dard length (SL), except for parts of the head, whichare expressed as percentages of head length. Countsof pectoral and caudal fin rays and vertebrae weremade only on cleared and stained specimens (c&s)prepared in accordance with Taylor and V an Dyke

(1985); in vertebral counts, the compound caudal cen-trum was counted as a single element. Selected osteo-logical features presented in the description are thosewith some relevance for clarifying phylogenetic rela-tionships among species of Rivulus, as described anddiscussed by Costa (1998). Nomenclature for frontalsquamation patterns and identification of homologousscales follows Hoedeman (1958); terminology forcephalic neuromasts follows Costa (2001). Compara-tive material is listed in Costa (1995a). Institutionalabbreviations are: MCP, Museu de Ciências e Tecnolo-gia da Pontifícia Universidade Católica do Rio Grandedo Sul, Porto Alegre, and UFRJ, Universidade Federaldo Rio de Janeiro, Rio de Janeiro.

Rivulus paracatuensis n. sp.(Fig. 2; Table I)

Rivulus decoratus non R. decoratus Costa, 1989:Costa, 1995a: 221-223 (misidentification).

Holotype: MCP 29639, male, 22.5 mm SL; Brazil:Estado de Minas Gerais, stream in Rio Paracatú flood-plains near Brasilândia de Minas, road MG 181, RioSão F rancisco b asin, a pproximately 1 7º00’00”S46º00’00”W; collected by Gilberto C. Brasil, 1 May 1994.Paratypes: MCP 29640, one female, 19.7 mm SL;UFRJ 2290, two males, 19.0-19.9 mm SL, and onefemale, 21.1 mm SL; UFRJ 2291, one male 23.9 mmSL, and one female 19.4 mm SL (c&s); same data asholotype.

DiagnosisSimilar to R. zygonectes Myers, R. punctatus

Boulenger, R. violaceus Costa, R. modestus Costa,R. pictus Costa, and R. decoratus Costa, and distin-guished from all other species of the genus by unpairedfins of female with contrasting dark grey marks and


Fig. 2. Rivulus paracatuensis, MCP 29637, male, holotype, 22.5 mm SL; Brazil: Minas Gerais: Brasilândia de Minas.Photo by W. J. E. M. Costa.

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black margins with white to yellowish white groundcolour (vs. never a similar colour pattern), dark brownto black postorbital stripe on head midline (vs. postor-bital stripe absent or present below head midline), andfirst epibranchial L-shaped (vs. I-shaped). Similar to R.pictus, but distinguished from R. zygonectes, R. punc-tatus, R. violaceus, R. modestus, and R. decoratus, bycaudal fin of male yellow with dark bars, without distinctcolours on edges (vs. having white edge in R. puncta-tus, distinctive yellow stripe on dorsal and ventral margins in R. zygonectes and R. violaceus , no mark-ings in R. modestus, and dark brown stripe on ventraledge in R. decoratus). Differs from R. pictus by usuallyhaving frontal squamation D-patterned, sometimes E-patterned (vs. usually F , sometimes E), side of bodypale blue in male (vs. bright greenish blue), and dorsaland caudal fins of male with dark grey bars (vs. red).Further distinguished from R. decoratus, the only con-gener also occurring in the São Francisco basin, bymore dorsal fin rays (9-10 vs. 7-8), more caudal fin rays(31-33 vs. 26-27), more pectoral fin rays (13 vs. 11-12),more vertebrae (30 vs. 28-29), more scales in longitu-dinal series (31-33 vs. 26-28), more scale rows aroundcaudal peduncle (16 vs. 14), and wider basihyal (widthabout 60% of longitudinal length, vs. about 35%).

DescriptionMorphometric data for holotype and four paratypes

are given in Table I. Male larger than female, largestmale 23.9 mm SL. Dorsal profile slightly convexbetween snout and end of dorsal fin base, gently con-

cave on caudal peduncle. Ventral profile slightly con-vex from lower jaw to end of anal fin base, nearlystraight on caudal peduncle. Body slender , subcylin-drical, body depth slightly greater than body width.Greatest body depth on vertical through pelvic finbase. Jaws short, snout blunt.

Tip of dorsal and anal fins rounded. Caudal fin ellip-tical. Pectoral fin elliptical, short, its posterior marginextending to vertical anterior to pelvic fin base. Pelvicfin short, its tip reaching base of first anal fin ray inmale, and anus in female. Dorsal fin origin on verticalbetween base of 9 th and 10th anal fin rays. Dorsal finrays 9-10, anal fin rays 13-14, caudal fin rays 31-33,pelvic fin rays 6-7, pectoral fin rays 13.

Scales large, cycloid. Body and head entirely scaled,except on chin. Frontal squamation usually D-pat-terned, sometimes E-patterned, scales arranged cir-cularly around central A-scale. No scales on dorsaland anal fin bases. Body squamation extendingslightly onto caudal fin base. Longitudinal series ofscales 31-33, transverse series of scales 8, scalerows around caudal peduncle 16. Contact organsabsent. Supraorbital neuromasts 3 + 3.

Ventral process of angulo-articular narrow and short.Dorsal arm of preopercle short and pointed. Basihyalapproximately triangular, its greatest width about 60 %of its total longitudinal length; basihyal cartilage short,about 20% of its total longitudinal length. Six bran-chiostegal rays. No teeth on second pharyngo-branchial. Deep gap lateral to anterior condyle of sec-ond pharyngobranchial. Distal region of first epi-


Males Females

Holotype Paratypes

MCP UFRJ UFRJ UFRJ MCP29637 2290 2290 2290 29638

SL [mm] 22.5 19.9 19.0 21.1 19.7

In percents of standard lengthBody depth 21.9 23.3 22.2 21.0 21.3Caudal peduncle depth 13.8 14.8 14.1 13.0 13.3Predorsal length 75.0 76.9 78.0 81.1 77.8Prepelvic length 53.3 55.9 55.3 56.7 57.1Length of dorsal fin base 12.6 12.8 11.8 9.3 10.2Length of anal fin base 22.2 20.0 20.0 18.1 17.9Caudal fin length 36.3 - 38.4 35.1 35.2Pectoral fin length 21.1 20.6 21.2 18.8 18.9Pelvic fin length 11.7 12.6 11.9 8.5 10.1Head length 26.7 28.2 28.2 27.5 27.6Head depth 19.0 20.2 18.3 18.1 18.5Head width 19.8 20.9 20.1 20.0 20.5

In percents of head lengthSnout length 15.4 15.7 14.4 15.3 14.0Lower jaw length 23.2 20.1 17.4 19.8 17.3Eye diameter 33.7 35.5 32.1 32.4 33.9

Table I. Morphometric data of Rivulus paracatuensis n. sp.

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branchial bent at angle of 90º to its main axis. Teethalong proximal and median portions of dorsal surfaceof fourth ceratobranchial. Gill-rakers of first branchialarch 1 + 7. Vomerine teeth 3-4. Interhyal not ossified.Posttemporal and supracleithrum fused. V entralprocess of posttemporal absent. Dorsal and ventralhypural plates separated by interspace. Total verte-brae 30.

Coloration in life: Male: side of body light yellowishbrown with faint light blue iridescence; stripe alongmidline, between head and anterior portion of caudalpeduncle, dark brown in its anteriormost part, becom-ing pale grey posteriorly; oblique rows of red dotsalong flank, forming chevron-like marks, with vertexdirected anteriorly, on posterior portion of flank. Headlight yellowish brown, opercular region with faint lightblue iridescence; lower jaw dark grey; dark brownstripe between point just posterior to posterior orbitaledge and humeral region, continuous with lateralstripe on body. Iris yellow. Dorsal and caudal fins yel-low with grey bars. Anal fin yellow, base light blue withred bars. Pectoral fin hyaline. Pelvic fin yellow .

Female: side of body light yellowish brown with mid-lateral stripe as in male, and oblique rows of reddishbrown dots. Head light yellowish brown, lower jawdark grey, and dark brown stripe posterior to orbit asin male. Iris yellow. Unpaired fins yellowish white withgrey bars; distal border of dorsal and anal fins, andentire border of caudal fin dark grey; black spot onupper portion of caudal fin base. Pectoral fin hyaline.Pelvic fin yellowish white.

EtymologyThe name paracatuensis refers to the type locality of

the new species, the rio Paracatú floodplains.

DistributionKnown only from the type locality, the floodplains of the

Rio Paracatú, Rio São Francisco basin, Brazil (Fig. 1).

DiscussionRivulus paracatuens is exhibits synapomorp hies

established by Costa (1995a, 1998) to define a cladewhich includes R. zygonectes , R. punctatus , R. vio-laceus, R. modestus , R. pictus , and R. decoratus :unpaired fins of female with dark grey marks andblack margins strongly contrasting with white to yel-lowish white background, dark brown to black postor-bital stripe on head midline, and distal portion of firstepibranchial bent at angle of 90º. This species assem-blage is endemic to central South America (Costa,1995a), which includes south-eastern Amazonian trib-utaries (R. modestus, R. violaceus, and R. zygo-nectes), Paraguay and Uruguay basins (R. puncta-tus), Paraná basin (R. pictus) and São Franciscobasin (R. decoratus and R. paracatuensis). Relation-ships within this clade are unclear at the presenttime.

AcknowledgementsSpecial thanks are due to Gilberto Brasil for collecting

and making available specimens included in the typeseries. This study was supported by CNPq-MCT (Con-selho Nacional de Desenvolvimento Científico e Tec-nológico – Ministério de Ciência e Tecnologia) andFAPERJ (Fundação de Amparo à Pesquisa do Estadodo Rio de Janeiro). The manuscript benefited fromrevision by M. Bailey.

ReferencesBritski, H. A., Y. Sato & A. B. S. Rosa. 1988. Manual

de identificação de peixes da região de T rês Marias(com chaves de identificação para os peixes da baciado São Francisco) (3ª edição). Ministério da Irrigação& CODEVASF, Brasília, 115 pp.

Costa, W. J. E. M. 1995a. Revision of the Rivuluspunctatus species-complex (Cyprinodontiformes:Rivulidae). Ichthyological Exploration of Freshwaters ,6(3): 207-226.

Costa, W. J. E. M. 1995b. Pearl killifishes, theCynolebiatinae: systematics and biogeography of aneotropical annual fish subfamily (Cyprinodonti-formes: Rivulidae). TFH, Neptune City, NJ, USA, 128pp.

Costa, W. J. E. M. 1998. Phylogeny and classificationof Rivulidae revisited: evolution of annualism andminiaturization in rivulid fishes (Cyprinodontiformes:Aplocheiloidei). Journal of Comparative Biology,3(1): 33-92.

Costa, W. J. E. M. 1989. Descrição de cinco novasespécies de Rivulus das bacias dos rios Paraná eSão Francisco (Cyprinodontiformes, Rivulidae).Revista Brasileira de Zoologia, 6(3): 523-534.

Costa, W. J. E. M. 2001. The neotropical annual fishgenus Cynolebias (Cyprinodontiformes: Rivulidae):phylogenetic relationships, taxonomic revision andbiogeography. Ichthyological Exploration of Freshwa-ters, 12(4): 333-383.

Costa, W. J. E. M. 2003. The Simpsonichthys flavicau-datus species group (Cyprinodontiformes: Rivulidae:Cynolebiatinae): phylogenetic relationships, taxo-nomic revision and biogeography. IchthyologicalExploration of Freshwaters, 14(1): 31-60.

Hoedeman, J. J. 1958. Rivulid fishes of the Antilles.Studies on the Fauna of Curaçao and otherCaribbean Islands, 32: 112-127.

Taylor, W. R. & G. C. Van Dyke. 1985. Revised proce-dures for staining and clearing small fishes and othervertebrates for bone and cartilage study . Cybium,9: 107-109.

Travassos, H. 1960. Catálogo dos peixes do vale dorio São Francisco. Boletim da

Sociedade Cearense de Agronomia, 1: 1-66.


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Index of aqua Vol. 6 (1-4)(Index by: 1. Author(s); 2. New Taxa; 3. Biology/Ecology/Biography/Reviews)

Author(s):Acero, Arturo P. and Betancur-R, Ricardo: Description of Arius neogranatensis, a new species of sea catfish from Colombia, with an identification key for Carribean ariid fishes. aqua 6 (1) 5-10, October 2002.Allen, Gerald R. and Bailey, Steven: Chrysiptera albata, a new species of damselfish (Pomacentridae) from the Phoenix Island, Central Pacific Ocean. aqua 6 (1) 39-43, October 2002.Allen, Gerald R.: Descriptions of two new species of damselfishes (Pomacentridae: Pomacentrus) from Madagascar. aqua 6 (2) 45-52, November 2002.Collette, Bruce B., Williams, Jeffrey T., Thacker, Christine E. and Smith, Michael L.: Shore fishes of Navassa Island, West Indies: a case study on the need for rotenone sampling in reef fish biodiversity studies. aqua 6 (3) 89-131, February 2003.Costa, Wilson J. E. M. and Cheffe, Morevy M.: Austrolebias jaegari (Cyprinodontiformes: Rivulidae: Cynolebiatinae): a new annual fish from the Laguna dosPatos system, southern Brazil, with a redescription of A. gymnoventris (Amato). aqua 6 (2) 83-88, November 2002.Costa, Wilson J. E. M., Moreira, Cristiano R. and Lima, Flávio C. T.: Simpsonichthys cholopteryx n. sp. (Cyprinodontiformes: Rivulidae: Cynolebiatinae): a new dwarf annual fish from the upper Rio Araguaia basin, central Brazil. aqua 6 (4) 139-144, March 2003.Dias, Thelma L. P. and Rosa, Ierecê L.: Habitat preferences of a seahorse species, Hippocampus reidi (Teleostei: Syngnathidae) in Brazil. aqua 6 (4) 165-176,March 2003.Feitoza, Bertran M.: Platygillellus brasiliensis n. sp. (Perciformes: Dactyloscopidae), the third species of the genus from the Atlantic. aqua 6 (1) 21-28, October2002.Gill, Anthony C. and Senou, Hiroshi: Lubbockchthys tanakai, new species of Pseudoplesiopine dottyback from the W est Pacific (Perciformes: Pseudochromidae).aqua 6 (1) 1-4, October 2002.Machado, Leonardo Francisco, Andrade, Áthila Bertoncini, Hostim–Silva, Maurício and Barreiros, João Pedro: Habitat use by the juvenile dusky grouperEpinephelus marginatus and its relative abundance, in Santa Catarina, Brazil. aqua 6 (4) 133-138, March 2003.Miquelarena, Amalia M., Protogino, Lucila C., Filiberto, Ramiro and López, Hugo L.: A new species of Bryconamericus (Characiformes: Characidae) from theCuña-Pirú creek in north-eastern Argentina, with comments on accompanying fishes. aqua 6 (2) 69-82, November 2002.Orlov, Alexei M.: Summer diet and feeding of shortraker (Sebastes borealis) and rougheye (S. aleutianus) rockfishes (Scorpaenidae) in the western Bering Sea.aqua 6 (1) 29-38, October 2002.Sazima, Ivan: Juvenile grunt (Haemulidae) mimicking a venomous leatherjacket (Carangidae), with a summary of Batesian mimicry in marine fishe s. aqua 6 (2) 61-68, November 2002.Soares, Marta S. C., Barreiros, João Pedro, Sousa, Luis and Santos, Ricardo S.: Agonistic and predatory behaviour of the lizardfish Synodus saurus(Linnaeus, 1758) (Actinopterygii: Synodontidae) from the Azores. aqua 6 (2) 53-60, November 2002.Vidthayanon, Chavalit and Jaruthanin, Kittipongse: Schistura kaysonei (Teleostei:Balitoridae), a new cave fish from the Khammouan karst, Laos PDR. aqua 6 (1) 17-20, October 2002.Williams, Jeffrey T.: Three new species of blennioid shore fishes discovered at Navassa Island, Carribean Sea. aqua 6 (1) 11-16, October 2002.Williams, Jeffrey T. and Mounts, Julie H.: Descriptions of six new Caribbean fish species in the genus Starksia (Labrisomidae). aqua 6 (4) 145-164, March 2003.

New Taxa:Acanthemblemaria harpeza n. sp.Three new species of blennioid shore fishes discovered at Navassa Island, Carribean Sea. aqua 6 (1) 11-16, October 2002.Arius neogranatensis n. sp. Description a new species of sea catfish from Colombia, with an identification key for Carribean ariid fishes. aqua 6 (1) 5-10, October 2002.Austrolebias jaegari n. sp. A new annual fish from the Laguna dos Patos system, southern Brazil, with a redescription of A. gymnoventris (Amato). aqua 6 (2) 83-88, November 2002.Bryconamericus menni n. sp. A new species of Bryconamericus (Characiformes: Characidae) from the Cuña-Pirú creek in north-eastern Argentina, with comments on accompanying fishes. aqua 6 (2) 69-82, November 2002.Chrysiptera albata n. sp. A new species of damselfish (Pomacentridae) from the Phoenix Island, Central Pacific Ocean. aqua 6 (1) 39-43, October 2002.Emblemaria vitta n. sp. Three new species of blennioid shore fishes discovered at Navassa Island, Carribean Sea. aqua 6 (1) 11-16, October 2002.Gillellus inescatus n. sp.Three new species of blennioid shore fishes discovered at Navassa Island, Carribean Sea. aqua 6 (1) 11-16, October 2002.Lubbockchthys tanakai n. sp. A new species of Pseudoplesiopine dottyback from the W est Pacific (Perciformes: Pseudochromidae). aqua 6 (1) 1-4, October2002.Platygillellus brasiliensis n. sp. The third species of the genus from the Atlantic. aqua 6 (1) 21-28, October 2002.Pomacentrus atriaxillaris n. sp. Descriptions of two new species of damselfishes (Pomacentridae: Pomacentrus) from Madagascar. aqua 6 (2) 45-52, November2002.Pomacentrus caeruleopunctatus n. sp. Descriptions of two new species of damselfishes (Pomacentridae: Pomacentrus) from Madagascar. aqua 6 (2) 45-52,November 2002.Schistura kaysonei n. sp. A new cave fish from the Khammouan karst, Laos PDR. aqua 6 (1) 17-20, October 2002.Simpsonichthys cholopteryx n. sp. A new dwarf annual fish from the upper Rio Araguaia basin, central Brazil. aqua 6 (4) 139-144, March 2003.Starksia leucovitta n. sp. Descriptions of six new Caribbean fish species in the genus Starksia (Labrisomidae). aqua 6 (4) 145-164, March 2003.Starksia melasma n. sp. Descriptions of six new Caribbean fish species in the genus Starksia (Labrisomidae). aqua 6 (4) 145-164, March 2003.Starksia multilepis n. sp. Descriptions of six new Caribbean fish species in the genus Starksia (Labrisomidae). aqua 6 (4) 145-164, March 2003.Starksia rava n. sp. Descriptions of six new Caribbean fish species in the genus Starksia (Labrisomidae). aqua 6 (4) 145-164, March 2003.Starksia sella n. sp. Descriptions of six new Caribbean fish species in the genus Starksia (Labrisomidae). aqua 6 (4) 145-164, March 2003.Starksia smithvanizi n. sp. Descriptions of six new Caribbean fish species in the genus Starksia (Labrisomidae). aqua 6 (4) 145-164, March 2003.

Biology/Ecology/Biography/Reviews:Agonistic and predatory behaviour of the lizardfish Synodus saurus (Linnaeus, 1758) (Actinopterygii: Synodontidae) from the Azores. aqua 6 (2) 53-60,November 2002.A new species of Bryconamericus (Characiformes: Characidae) from the Cuña-Pirú creek in north-eastern Argentina, with comments on accompanyingfishes. aqua 6 (2) 69-82, November 2002.Austrolebias jaegari (Cyprinodontiformes: Rivulidae: Cynolebiatinae): a new annual fish from the Laguna dos Patos system, southern Brazil, with aredescription of A. gymnoventris (Amato). aqua 6 (2) 83-88, November 2002.Book review: Bloch’s fish collection in the Museum für Naturkunde der Humboldt Universität zu Berlin – an illustrated catalogue and historic al accountand Block’s Atlas – anniversary edition. aqua 6 (3) 132, February 2003.Habitat preferences of a seahorse species, Hippocampus reidi (Teleostei: Syngnathidae) in Brazil. aqua 6 (4) 165-176, March 2003.Habitat use by the juvenile dusky grouper Epinephelus marginatus and its relative abundance, in Santa Catarina, Brazil. aqua 6 (4) 133-138, March 2003.Juvenile grunt (Haemulidae) mimicking a venomous leatherjacket (Carangidae), with a summary of Batesian mimicry in marine fishe s. aqua 6 (2) 61-68,November 2002.Shore fishes of Navassa Island, West Indies: a case study on the need for rotenone sampling in reef fish biodiversity studies. aqua 6 (3) 89-131, February 2003.Summer diet and feeding of shortraker (Sebastes borealis) and rougheye (S. aleutianus) rockfishes (Scorpaenidae) in the western Bering Sea.aqua 6 (1) 29-38, October 2002.

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Instructions to Authorsaqua is an international journal which publishes original sci-

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Day, J. H., Blaber , S. J. M., & J. H. W allace. 1981. EstuarineFishes. In: Estuarine Ecology with Particular Reference to South-ern Africa. (Ed. J.H. Day .) : 197-221. A. A. Balkema, Rotterdam.

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aquaJournal of Ichthyology and Aquatic Biology

Vol. 7 (1), May 2003

Contents

John E. Randall: Thalassoma nigrofasciatum, a new species of labrid fish from the south-west Pacific ............................................................................................................................................1-8

Richard Winterbottom: A new species of Trimma (Gobiidae) from the western Indian Ocean........................................................................................................................................................9-12

Richard Winterbottom and Cesar A. Villa: A new species of the Trimma caesiura complex (Gobiidae, Teleostei) from the north–eastern margin of the Australian Plate, with a redescription of the other nominal species in the complex .....................................................................13-28

Marta S. C. Soares, Luis Sousa and João Pedro Barreiros: Feeding habits of the lizardfish Synodus saurus (Linnaeus, 1758) (Actinopterygii: Synodontidae) from the Azores..................................................................................................................................................29-38

Wilson J. E. M. Costa: Rivulus paracatuensis n. sp. (Cyprinodontiformes: Rivulidae): a new rivuline species from the Rio São Francisco basin, Brazil ......................................................................39-43

Index of aqua Vol. 6 (1-4) .......................................................................................................................................44

Papers appearing in this journal are indexed in: Zoological Record; Biolis - Biologische Literatur Information Senckenberg;

www.aquageo.com; www.Joachim-Frische.com

Cover photo: Trimma caesiura, live, Palau. Photo by H. Nagano.

A bay among the Belmont, Challenger and Cabral Islets. Photo by B. M. Feitoza from a forthcoming MS in aqua 7(2).

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7(1)imp/ok 16-06-2003 11:33 pagina 1 aqua1).pdf · thalassoma nigrofasciatum est décrit en tant...

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