International Journal of Ichthyology3).pdf · International Journal of Ichthyology Vol. 17 (3), 10 July 2011 Aquapress ISSN 0945-9871. Managing Editor: Heiko Bleher Via G. Falcone

aquaInternational Journal of Ichthyology

Vol. 17 (3), 10 July 2011

AquapressISSN

0945-9871

Managing Editor:

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

Scientific Editor:

Helen Larson65 Reid Road, Wongaling Beach, Queensland 4852, AustraliaTel: +61-7-4068.8095E-mail: [email protected]

Editorial Board:Gerald R. Allen Department of Aquatic Zoology, Western Australian Museum, Perth, Australia

Nina G. Bogutskaya Zoological Institute of the Russian Academy of Sciences, St. Petersburg, Russia

Friedhelm KruppCurator of Fishes, Senckenberg Research Institute andNatural History Museum, Frankfurt am Main, Germany

Flávio C. T. LimaMuseu de Zoologia da Universidade de São PauloSão Paulo, Brasil

Axel Meyer Lehrstuhl für Zoologie und Evolutions biologie, Universität Konstanz, Germany

Paolo Parenti Department of Enviromental Sciences, University of Milano-Bicocca, Milan, Italy

Mário de PinnaMuseu de Zoologia da USP, São Paulo, Brazil

John E. RandallBishop Museum, Honolulu, Hawaii, U.S.A.

Richard WinterbottomCentre of Biodiversity & Conservation Biology, Royal Ontario Museum, Toronto, Canada

Scopeaqua is an international journal which publishes originalscientific articles in the fields of systematics, taxonomy,bio geography, ethology, ecology, and general biology offishes. Papers on freshwater, brackish, and marine fisheswill be considered. aqua is fully refereed and aims at pub-lishing manuscripts within 2-4 months of acceptance. Inview of the importance of color patterns in species identi -fication and animal ethology, authors are encouraged tosubmit color illustrations in addition to descriptions ofcoloration. It is our aim to provide the international sci-entific community with an efficiently published journalmeeting high scientific and technical standards.

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ISSN 0945-9871Publisher: Aquapress, Redazione aqua, I-27010 Miradolo Terme (Pavia), Italywww.aqua-aquapress.comPrinter: Global Print s.r.l. – Gorgonzola (MI) – ItalyCopyediting and layout: Rossella Bulla© 2011 aqua, International Journal of Ichthyology

aqua - International Journal of Ichthyology

AbstractStudies of the feeding habits of reef-inhabiting herbivo-

rous species are important since they are the first level inthe trophic chain between primary producers and con-sumers. This study is the first contribution to determinethe diet of the convict surgeonfish Acanthurus triostegusfrom the Los Frailes reef, Baja California Sur, Mexico. Wedescribe qualitative and quantitatively the trophic spec-trum of 50 A. triostegus individuals, using the traditionalmethods of frequency of occurrence and weight, as well asthe index of relative importance. Overall, we identified 35food components, of which 18 were algae of the ClassRhodophyceae, 10 of the Chlorophyceae and six Phaeo-phyceae. We determined that A. triostegus is a strict diur-nal herbivore, which mainly feeds on the green alga Ulvalinza, and even though it is a species with a wide geo-graphic distribution, its behavioral feeding pattern ishomogeneous.

ZusammenfassungDie Untersuchung des Nahrungsverhaltens pflanzenfres -

sen der Arten unter den Riffbewohnern hat eine besondereBedeutung, weil sie in der Nahrungskette zwischenPrimärproduzenten und Konsumenten die erste Stufedarstellen. In der vorliegenden Arbeit wird erstmals dieErnährung des Sträflings-Doktorfisches Acanthurus trioste-gus beispielhaft an Tieren untersucht, die über dem Frailes-Riff in Baja California Sur, Mexiko, leben. Wirbeschreiben das qualitative und quantitativeNahrungsspektrum von 50 Individuen der Art A. triostegusnach den üblichen Methoden durch Häufigkeit undGewicht der Nahrungsbestandteile sowie den Index der re -lativen Bedeutung. Insgesamt ermittelten wir 35 Nah -rungs bestandteile, darunter gehörten 18 Algenarten derKlasse Rhodophyceae an, 10 den Chlorophyceae und sechsden Phaeophyceae. Wir konnten feststellen, dass A. trioste-gus streng tagaktiv herbivor lebt und sich hauptsächlichvon der Grünalge Ulva linza ernährt; trotz ihrer weitengeografischen Verbreitung ist das Nahrungsverhaltendieser Doktorfisch-Art homogen.

RésuméLes études portant sur les habitudes alimentaires d’e-

spèces herbivores récifales sont importantes dans la mesureoù elles forment le premier échelon de la chaine alimen-taire entre les producteurs primaires et les consommateurs.Cette étude est la première à analyser le régime duChirurgien Acanthurus triostegus du récif de Los Frailes,Baja California Sur, Mexique. Nous présentons le spectretrophique qualitatif et quantitatif de 50 A. triostegus àl’aide des méthodes traditionnelles de nombre et de poidsainsi que de l’index de l’importance relative. En résumé,nous avons relevé 35 sortes d’aliments dont 19 étaient desalgues de la classe des Rhodophyceae, 10 des Chloro-phyceae et 6 des Phaeophyceae. Nous avons constaté qu’A.triostegus est un herbivore exclusivement diurne qui senourrit surtout de l’algue verte Ulva linza et, bien que l’espèce connaisse une large distribution géographique, soncomportement alimentaire est homogène.

SommarioGli studi sulle abitudini alimentari delle specie erbivore

che abitano le barriere coralline hanno il merito di illu -strare il primo livello della catena trofica tra produttori pri-mari e consumatori. Questo studio rappresenta il primocontributo che descrive la dieta del pesce chirurgo carce -rato Acanthurus triostegus di Los Frailes reef, Baja Califor-nia Sur, Messico. Si descrive lo spettro trofico qualitativoquantitativo di 50 individui di A. triostegus, usando i me -todi tradizionali di frequenza delle presenze e del peso, co -me pure l’indice di importanza relativa. Nel complesso,sono stati identificati 35 componenti alimentari, di cui 18erano alghe della classe delle Rhodophyceae, 10 di quelladelle Chlorophyceae e sei delle Phaeophyceae. Si è deter-minato che A. triostegus è un erbivoro strettamente diurno,che si nutre principalmente dell’alga verde Ulva linza eche, anche se è una specie con un’ampia distribuzione geo -grafica, ha un comportamento alimentare assolutamenteomogeneo.

aqua vol. 17 no. 3 - 10 July 2011121

aqua, International Journal of Ichthyology

Feeding habits of the convict surgeonfish Acanthurus triostegus(Teleostei: Acanthuridae) on the Los Frailes reef, Baja California Sur, Mexico

Leonardo A. Abitia-Cárdenas, Xchel G. Moreno-Sánchez*, Deivis S. Palacios-Salgado and Ofelia Escobar-Sánchez

Centro Interdisciplinario de Ciencias Marinas (CICIMAR-IPN), Departamento de Pesquerías y BiologíaMarina. Apdo. Postal 592. La Paz, Baja California Sur, México. C.P. 23000.

*E-mail: [email protected]

Received: 23 August 2010 – Accepted: 08 January 2011

INTRODUCTIONThe Acanthuridae or surgeonfish are distributed

in tropical and subtropical oceans. There are 80known species distributed in six genera (Nelson2006). The convict surgeonfish, Acanthurus trioste-gus (Linnaeus, 1758), is probably the species withwidest distribution, having populations from theIndian Ocean to the eastern tropical Pacific. Thewide geographic range of this species has allowedthe differentiation of populations. Randall (1956)recognized three subspecies: Acanthurus triostegussandvicensis for Hawaii, A. triostegus marquesensisfor the Marquesas Islands in the central Pacific andAcanthurus t. triostegus for the rest of its distribu-tion, including the Line Islands and eastern tropi-cal Pacific. Genetic studies have shown strong dif-ferences among populations, with a high geneticdiversity for the Hawaiian populations, which val-idates the subspecies name sandvicensis Randall,1956, for the convict surgeonfish in that area(Planes & Fauvelot 2002).The longevity of convict surgeonfish individuals

is of about 40 years (Longenecker et al. 2008); theyhave separate sexes and reproduce during the dayin schools of 200 to 2000 individuals (Craig1998). Females mature (164 mm; 440 days) laterthan males (90 mm; 168 days) and reach greatersizes. There are estimates of up to 362 000 eggs peryear (Longenecker et al. 2008), and the larvalphase can last 44 to 60 days, which favors a widedistribution (McCormick 1999).In the Gulf of California there are six species from

two acanthurid genera, most with an Indo-Pacificaffinity (Robertson & Allen 2002), among them theconvict surgeonfish A. triostegus. In particular in theLos Frailes reef area, which is within the perimeter ofthe Marine Park of Cabo Pulmo in Baja CaliforniaSur (B.C.S.), Mexico, A. triostegus is a species con-sidered to be common and is among the 12 mostdominant species on the reef (Moreno 2009).However, there is no information on the trophic

biology of the species in the area. The few studieson trophic ecology of herbivorous fish in the Gulfof California have been carried out on indigenousspecies that cohabit with A. triostegus, such as theyellowtail surgeonfish Prionorus punctatus (Mont-gomery et al. 1980), the giant damselfishMicrospathodon dorsalis and the Cortez damselfishStegastes rectifraenum (Montgomery 1980a, b).However, there are studies carried out in the barrierreef of Aldabra, Indian Ocean (Robertson &Gaines 1986) and in the Hawaiian Islands (Randall

1961), where the trophic spectrum of A. triosteguswas characterized.In the present study we characterize the diet of A.

triostegus and consider the diversity, frequency andabundance of algae species consumed by this fish.

MATERIAL AND METHODSThe fish were captured monthly from November

2004 to October 2005 in the rocky reef of LosFrailes (23°5’ N and 109°5’ W), in the southernpart of the Baja California peninsula, Mexico.Specimens were captured using a pole spear by afree diver between 10:00 and 16:00 hours, the timeof best visibility. For each collected fish the totalweight (TW) and total length (TL) were recordedand the stomach was extracted. Gastric contentswere put in plastic bags and frozen for later analy-sis at the Fish Ecology laboratory at the CentroInterdisciplinario de Ciencias Marinas (CICI-MAR-IPN) in La Paz, Baja California Sur.During stomach content analysis we separated

food categories (items) according to taxonomicgroup, identifying each to the minimum possibletaxon, depending on digestion stage. For the taxo-nomic identification of algae we used the keys byDawson (1944, 1961), Abbott & Hollenberg(1976) and Espinoza-Avalos (1993).Once the taxonomic work was completed, we

analyzed stomach contents quantitatively, forwhich we used: The percentage of frequency of occurrence

(%FO), referred as the frequency of occurrence ofprey items within the total number of stomachswith food.%FO= No. of stomachs including a prey item-

No. of stomachs with food × 100The gravimetric composition percentage (%W) is

the wet weight of prey items found within the totalwet weight of stomachs with food:%W= Weight of prey item- Total weight of prey

items × 100It is worth noting that due to the nature of the

feeding habits of this species (algae grazers) it wasnot possible to employ a numeric method to quan-tify units. In order to evaluate in an integral man-ner the importance of each food type in thespecies´ diet, we used the Index of Relative Impor-tance (IRI) of Pinkas et al. (1971), modified byOjeda & Muñoz (1999):IRI = % FO × %WWe calculated the diet width (Bi) using the stan-

dardized Levin´s index (Hurlbert 1978) from the

aqua vol. 17 no. 3 - 10 July 2011 122

Feeding habits of the convict surgeonfish Acanthurus triostegus (Teleostei: Acanthuridae) on the Los Frailes reef, Baja California Sur, Mexico

absolute values obtained using the gravimetricmethod. This index takes values from 0 to 1. WhenBi values are under 0.6, for example, a predator isconsidered a specialist, which indicates that it usesa low number of resources and presents a prefer-ence for certain food items. When values are closerto one (0.6) the spectrum is of a generalist, mean-ing that species uses all resources without selection.

RESULTSOverall we collected 50 fish (all had stomach con-

tents). We identified 35 items, of which 18 werealgae from the Class Rhodophyceae, 10 wereChlorophyceae and six were Phaeophyceae (Table I).

Using the gravimetric method, all items had atotal biomass of 225.08 g. The green alga Ulvalinza represented 42.08% (94.71 g), non-identi-fied organic matter (NIOM) represented 13.79%(31.04 g), Gelidiella spp. represented 12.36%(27.83 g), Polysiphonia simplex represented 7.32%(16.47 g) and Porphyra spp. represented 3.36%(7.56 g) (Fig. 1).Using the frequency of occurrence method, the

green alga Ulva linza was recorded in 56% of stom-achs (28 stomachs) followed by Gelidiella and P.simplex with 54% (27 stomachs) respectively, whilethe red algae Amphiroa valonioides and Laurenciawere recorded in 44% of stomachs (22 stomachs),

aqua vol. 17 no. 3 - 10 July 2011123

Leonardo A. Abitia-Cárdenas, Xchel G. Moreno-Sánchez, Deivis S. Palacios-Salgado and Ofelia Escobar-Sánchez

Class Species W %W FO %FO IRI %IRI

Chlorophyceae Ulva linza 94.7 42.1 28.0 56.0 2356.4 51.50Cladophora spp. 3.44 1.53 16.00 32.00 48.94 1.07Derbesia marina 2.11 0.94 7.00 14.00 13.11 0.29Ulva lactuca 1.99 0.88 11.00 22.00 19.41 0.42Bryopsis spp. 1.78 0.79 18.00 36.00 28.53 0.62Rhizoclonium spp. 1.68 0.75 9.00 18.00 13.43 0.29Codium simulans 0.19 0.08 3.00 6.00 0.50 0.01Caulerpa racemosa 0.15 0.06 1.00 2.00 0.13 0.00Cladophorosis fasciculatus 0.09 0.04 1.00 2.00 0.08 0.00Enteromorpha spp. 0.08 0.03 1.00 2.00 0.07 0.00

Rhodophyceae Gelidiella spp. 27.83 12.36 27.00 54.00 667.68 14.59Polysiphonia simplex 16.47 7.32 27.00 54.00 395.12 8.64Porphyra spp. 7.56 3.36 7.00 14.00 47.05 1.03Amphiroa valonioides 6.12 2.72 22.00 44.00 119.54 2.61Laurencia spp. 5.92 2.63 22.00 44.00 115.72 2.53Hypnea musciformis 3.58 1.59 12.00 24.00 38.19 0.84Champia spp. 3.30 1.47 14.00 28.00 41.04 0.90Jania mexicana 3.28 1.46 11.00 22.00 32.08 0.70Gracilaria spp. 2.26 1.00 8.00 16.00 16.06 0.35Herposiphonia spp. 1.68 0.75 4.00 8.00 5.98 0.13Pitophilium spp. 0.83 0.37 1.00 2.00 0.74 0.02Ceramium spp. 0.48 0.21 3.00 6.00 1.29 0.03Pterocladiella spp. 0.38 0.17 3.00 6.00 1.03 0.02Ahnfeltia spp. 0.20 0.09 1.00 2.00 0.18 0.00Centroceras spp. 0.11 0.05 1.00 2.00 0.10 0.00Gracilaria velanoae 0.07 0.03 2.00 4.00 0.12 0.00Goniotrichum alsidi 0.07 0.03 1.00 2.00 0.06 0.00Erythrotrichia spp. 0.05 0.02 1.00 2.00 0.05 0.00

Phaeophyceae Dictyota crenulata 2.14 0.95 14.00 28.00 26.61 0.58Sphaelaria spp. 1.93 0.86 10.00 20.00 17.14 0.38Lobophora spp. 1.58 0.70 6.00 12.00 8.45 0.19Ectocarus spp. 1.22 0.54 6.00 12.00 6.52 0.14Dictyopteris delicatula 0.74 0.33 4.00 8.00 2.62 0.06Padina concrescens 0.01 0.01 1.00 2.00 0.01 0.00*NIOM 31.04 13.79 20.00 40.00 551.70 12.05

TOTAL 225 100 50 4576 100

Table I. Absolute and percent values of the frequency of occurrence (FO), gravimetric (W) and index of relative importance(IRI) methods of the Acanthurus triostegus diet in reef Los Frailes, B.C.S. * NIOM (non-identified organic matter).

Fig. 1. Main food compounds of Acanthurus triostegus, presented as percentage weight, frequency of occurrence and index ofrelative importance (%IRI). * NIOM (non-identified organic matter).


124aqua vol. 17 no. 3 - 10 July 2011

Likewise, Randall (1961) described the diet of A.triostegus sandvicensis captured in the HawaiianIslands, and stated that it fed preferentially on thered alga Polysiphonia and on the green alga Entero-morpha, completing its diet with a variety of algaeamong which the most important were Hypnea,Ceramium, Gracilaria and Rhizoclonium species.The dominance in the diet of the green alga Ulva

linza and the red algae Gelidiella and Polysiphoniasimplex, as well as the low Levin´s index value (Bi =0.10), indicated that A. triostegus has a specialisttrophic behavior. Among the factors that can deter-mine the food selection patterns by herbivorousfish are availability and relative abundance of thefood (Horn 1983), the hardness of the algal thal-lus, the assimilation efficiency and the nutritionalvalue (Littler & Littler 1980), as well as the pres-ence of secondary compounds present in algae(Targett & Targett 1990). It is interesting to notethat in the present study a secondary elementfound in the stomachs of A. triostegus was sand.This has been reported for other acanthurid speciesand it is thought that its presence can help the fishmore rapidly and efficiently grind up plant tissues,which would optimize digestion and energy assim-ilation (Randall 1967).The indigenous herbivorous fish species with

which Acanthurus triostegus cohabits in the Los

and NIOM was recorded in 40% of stomachs (20stomachs) (Fig. 1). According to the IRI, the mostimportant food item was Ulva linza, which repre-sented 51.50% of the A. triostegus diet, followed byGelidiella spp. with 14.59%, NIOM with 12.06%,and Polysiphonia simplex with 8.64%. The Levin´sindex showed that the diet width of A. triostegus issmall (Bi = 0.10), so that from its trophic behaviorit can be considered a specialist.

DISCUSSIONThe convict surgeonfish A. triostegus is a strict

herbivore, since all recorded items were algae, con-firming the report by Robertson & Allen (2002),who mentioned that acanthurids feed on algaethroughout tropical Pacific Ocean reefs. For theLos Frailes area in particular, we identified 35 algaespecies, which represents 55.7% of the flora diver-sity reported for the Cabo Pulmo-Los Frailes areaby Anaya-Reyna & Riosmena-Rodríguez (1996).Acanthurus triostegus in the Los Frailes reef fed on

a high proportion of green and red algae and on asmaller proportion of brown algae. Robertson &Gaines (1986) detected this same feeding patternfor the species at Aldabra Atoll in the IndianOcean. In that study the green alga Ulva was thefourth in importance, in contrast to the presentstudy in which Ulva linza was the most important.

Leonardo A. Abitia-Cárdenas, Xchel G. Moreno-Sánchez, Deivis S. Palacios-Salgado and Ofelia Escobar-Sánchez

125 aqua vol. 17 no. 3 - 10 July 2011

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Frailes reef feed on the same algal assemblage. Theyellowtail surgeonfish Prionurus punctatus con-sumes green and red algae (Ulva linza and Geli-diella) (Montgomery et al. 1980), the giant dam-selfish Microspathodon dorsalis is a non-selective her-bivore that consumes a large proportion of red algaePolysiphonia (60.5%), Gracilaria (8.2%) andCeramium (5.8%) while the Cortez damselfish Ste-gastes rectifraenum feeds on a wide variety of species,among them red algae such as Gracilaria (36.4%),Ceramium (7.1%) and Polysiphonia (7.2%), as wellas green algae Ulva (16.1%) (Montgomery 1980a,b). However, in most cases where a similarity indiets has been reported, the effect of high foodabundance can be seen, and when the resource isnot abundant other factors come into play such asspatial and temporal segregation, as well as trophicplasticity of the species that interact; therefore, it isconsidered that these species do not compete forfood (Moreno et al. 2009). Apart from alimentary segregation, such as was

shown in this study, fish can also segregate spatiallywithin the same reef. For example, acanthuridsform resident mobile schools with diurnal habits,with variable densities. Acanthurus triostegus formstemporal monospecific schools; Prionurus puncta-tus forms monospecific or mixed schools with P.laticlavius, mainly in shallow areas; while theschools of Acanthurus xanthopterus prefer the reefedges (Montgomery et al. 1980; Thomson et al.2000). The pomacentrids Microspathodon bairdii,M. dorsalis, Stegastes rectifraenum, S. acapulcoensisand S. flavilatus are territorial species that vigor-ously defend their refuge from intruders, includingconspecifics and other herbivorous fish (Welling-ton & Victor 1988), which allows the coexistenceof several herbivorous species in the area.According to the results obtained, we confirm

that Acanthurus triostegus is a diurnal herbivorousfish and a substrate grazer, which had already beendocumented for this species and other species fromthe same genus (A. chirurgus, A. bahianus and A.coeruleus) (Randall 1967).

ACKNOWLEDGEMENTSWe would like to thank the Instituto Politécnico

Nacional (IPN) for funds received through COFAAand EDI. Xchel G. Moreno Sánchez, Deivis S.Palacios Salgado and Ofelia Escobar Sánchez thankthe Consejo Nacional de Ciencia y Tecnología(CONACyT) and the Programa Institucional deFormación de Investigadores (PIFI-IPN).


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Six new species of the genus Trimma (Percomorpha; Gobiidae) from the RajaAmpat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature

Richard Winterbottom

Department of Natural History, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario, M5S 2C6; and Department of Ecology and Evolutionary Biology, University of Toronto,

Toronto, Ontario, M5S 3G5. Email: [email protected]

Received: 30 December 2010 – Accepted: 01 April 2011

127 aqua vol. 17 no. 3 - 10 July 2011

by a relatively deep body, numerous irregular scales withabout 30 lateral rows and over 15 scales in the anterior trans-verse series, a fifth pelvic fin ray that branches twice dichoto-mously, usually a somewhat elongate second dorsal spine, nopredorsal, cheek or opercular scales, a moderately wide bonyinterorbital with a fleshy median ridge between the eyes, anda dermal ridge anterior to the first dorsal spine. There is a redspot or elongate blotch above the opercle in live and freshmaterial, and the dorsal surface of the snout has a reticulateddark pattern, with dark transverse stripes over the dorsal mar-gin of the orbit. The species is known from south-westernSulawesi north to Palawan and eastwards to the SolomonIslands. Trimma papayum n. sp. is unique among thedescribed species of the genus in having a one-third pupildiameter black ocellated spot on and just behind the fourthdorsal fin spine. It has 9 dorsal and 8 anal fin rays, a singlebranch point in the fifth pelvic fin ray, 10-11 anterior and 8-9 posterior transverse scale rows, 5-10 scales in the predorsalmidline, and a single row of 3 scales on the upper margin ofthe opercle. Freshly collected specimens are orange red inoverall colouration, with scattered diffuse yellow spots. Thespecies is known only from Indonesia, at Maumere, Floresand Kawe Island, Raja Ampat. Trimma xanthochrum n. sp. ischaracterized by a wide interorbital region (80-100% pupildiameter), a second dorsal spine usually reaching posteriorlyto between the bases of the second to third dorsal fin rays, 15-16 pectoral rays usually with 7-8 branched rays, vertical rowsof sensory papillae below eye of 2-3 papillae in rows 1-4 and4-5 in row 5, a caudal blotch which has a lower half abouttwo-thirds the width of the upper half, and usually an overallyellowish body with yellow at least proximally in the caudalfin. It is currently known with certainty only from the RajaAmpat islands.

ZusammenfassungBei einer neueren Forschungsreise zum Raja-Ampat-Archi-

pel Indonesiens (2010) wurden sieben unbeschriebeneTrimma-Arten entdeckt, von denen sechs hier beschriebenwerden. Zwei der neuen Arten kommen nach heutigerKenntnis nur an diesen Inseln vor, die anderen sind auch vonanderen Teilen des Westpazifiks bekannt. Trimma cheni n. sp.unterscheidet sich von den anderen Trimma-Arten durch fol-

AbstractRecent (2010) fieldwork in the Raja Ampat Islands of

Indonesia resulted in the collection of seven undescribedspecies of Trimma, six of which are described here. Two ofthese are currently only known from these islands, the othersare known from other parts of the western Pacific. Trimmacheni n. sp. is distinguished from other species of Trimma inhaving 8-9 scales in the predorsal midline, 2-3 scales on theopercle, an elongate second dorsal spine reaching posteriorlyto the bases of rays 1-4 of the second dorsal fin, middle pec-toral fin rays branched, fifth pelvic fin ray branched once, adark basal stripe in the dorsal fins, scale pockets indistinctlyoutlined with darker pigment, and, in life, two red to orangebars across the cheek and three diffuse yellow stripes on thebody (most obvious along the caudal peduncle). It has beenrecorded from Palau, the Philippines, and Sulawesi and Flo-res in Indonesia. Trimma erdmanni n. sp. has a reddish-orange body with a dark red lateral stripe with darker borderson the body which extends anteriorly onto the head, where itbifurcates, and there is a thin longitudinal light stripe belowthe eye. It usually lacks scales in the predorsal midline, thesecond spine of the first dorsal is elongated, there are 9 dorsaland 8 anal rays, a single branch in the fifth pelvic fin ray, and19-22 gill rakers on the first gill arch. The species is knownfrom the Raja Ampat islands, the Hermit Islands andMadang (Papua New Guinea), and from photographs fromEl Nido, Palawan Island and Davao Gulf, Mindanoro,Philippines. Trimma habrum n. sp. has a bony interorbitalwidth equal to the pupil diameter, 8-9 scales in the predorsalmidline, 14 unbranched pectoral fin rays, an unbranchedfifth pelvic fin ray, no basal membrane joining the fifth pelvicfin rays across the midline, usually a single full row of cheekscales, and scales on the upper two-thirds of the opercle. Thefresh colouration is diagnostic: a pale translucent dorsumwith light yellow blotches and the base of each element of thedorsal fin surrounded by a red spot, a thin red bar along theposterior margins of the hypurals, no dark pigment at all onthe hypural region of the peduncle, and a darkly pigmentedcovering to the dorsal margins of the swim bladder, neuralsheath, and the brain. It is currently known only from a sin-gle collection made at Kerou Island, Fam Islands in RajaAmpat, Indonesia. Trimma haimassum n. sp. is characterized

gende Merkmale: 8-9 Schuppen auf der prädorsalen Mittel-linie, 2-3 Schuppen auf dem Kiemendeckel, einen verlänger-ten zweiten Rückenstachel, der nach hinten bis zur Basis derFlossenstrahlen 1-4 der zweiten Rückenflosse reicht, ver-zweigte mittlere Strahlen der Brustflosse, einmal verzweigtenfünften Bauchflossenstrahl, einen dunklen Streifen amGrund der Rückenflossen, undeutlich mit dunklerem Farb-stoff umrahmte Schuppentaschen, außerdem beim lebendenTier zwei rote bis orangefarbene Bänder über die Wange hin-weg und drei schwach gelbe Streifen auf dem Rumpf (amdeutlichsten sichtbar auf dem Schwanzstiel). Diese Artkonnte bei Palau, den Philippinen und Sulawesi und an derInsel Flores in Indonesien nachgewiesen werden. Trimma erd-manni n. sp. hat einen rötlich orangefarbenen Rumpf undeinen dunkelroten Seitenstreifen mit dunkleren Rändern aufdem Rumpf, der sich nach vorne bis zum Kopf erstreckt, woer sich gabelt, außerdem einen dünnen hellen Längsstreifenunter dem Auge. Normalerweise fehlen hier Schuppen aufder prädorsalen Mittellinie, der zweite Stachel der erstenRückenflosse ist verlängert, die Rückenflossen haben 9 Flos-senstrahlen, die Afterflosse 8, der fünfte Bauchflossenstrahlverzweigt sich einmal, und der erste Kiemenbogen trägt 19-22 Kiemenreusen. Fundorte dieser Art sind die Inseln RajaAmpat, die Hermit-Inseln und Madang (Papua-Neuguinea)sowie, durch Fotos belegt, El Nido, Palawan-Insel und derDavao-Golf, Mindanoro, Philippinen. Bei Trimma habrumn. sp. entspricht die Breite des Interorbitalknochens demPupillendurchmesser, Vertreter dieser Art haben 8-9 Schup-pen auf der prädorsalen Mittellinie, 14 unverzweigte Brust-flossenstrahlen, einen unverzweigten fünften Bauchflos-senstrahl, keine Basalmembran am fünften Bauchflos-senstrahl über die Mittellinie hinweg, normalerweise eine ein-zelne volle Reihe von Wangenschuppen und Schuppen aufdem oberen Zweidrittel des Kiemendeckels. Außerdem ist dieFärbung lebender oder frischtoter Exemplare artkennzeich-nend: der Rücken blass durchscheinend mit hellgelbenFlecken, die Basis von jedem Element der Rückenflosse voneinem roten Fleck umrahmt, ein dünner roter Streifen ent-lang den hinteren Rändern der Hypurale, überhaupt keindunkles Pigment in der hypuralen Gegend des Schwanzstielssowie eine dunkel pigmentierte Bedeckung der dorsalen Rän-der der Schwimmblase, der Nervenscheide und des Gehirns.Bisher ist diese neue Art nur von einem einzigen Forschungs-fang an der Kerou-Insel, Fam-Inseln im Raja-Ampat-Archi-pel, Indonesien, bekannt. Trimma haimassum n. sp. istgekennzeichnet durch einen relativ tiefen Rumpf, zahlreicheunregelmäßige Schuppen mit etwa 30 seitlichen Reihen undüber 15 Schuppen in der vorderen Querreihe, einen fünftenBauchflossenstrahl, der sich zweimal dichotom verzweigt,normalerweise einen etwas verlängerten zweiten Rückensta-chel, das Fehlen von Schuppen prädorsal, an der Wange undauf dem Kiemendeckel, einen mäßig breiten Interorbitalkno-chen mit einem fleischigen medianen Grat zwischen denAugen sowie eine Hautleiste vor dem ersten Rückenstachel.Bei lebenden oder frischtoten Exemplaren zeigt sich in roterFarbe ein Punkt oder länglicher Fleck oberhalb vom Kie-mendeckel, und die rückwärtige Oberfläche der Schnauzeweist ein netzartiges dunkles Muster auf, mit dunklen Quer-

streifen über dem rückwärtigen Rand der Augenhöhle. Manfand diese Art am südwestlichen Sulawesi nördlich von Pala-wan und nach Osten zu an den Salomoninseln. Trimmapapayum n. sp. zeichnet sich unter den beschriebenen Artendadurch aus, dass sich auf und direkt hinter dem viertenRückenflossenstachel ein schwarzer Augenfleck mit demDurchmesser eines Drittels der Pupille befindet. Außerdemhaben Vertreter dieser Art 9 Rückenflossen- und 8 Afterflos-senstrahlen, nur einen Verzweigungspunkt beim fünftenBauchflossenstrahl, 10-11 vordere und 8-9 hintere quer lau-fende Schuppenreihen, 5-10 Schuppen auf der prädorsalenMittellinie und eine einzelne Reihe von drei Schuppen amoberen Rand des Kiemendeckels. Frisch gefangene Tiere sindinsgesamt orangerot, mit unregelmäßig verteilten schwachgelben Flecken. Bisher ist die Art nur von Indonesienbekannt, von der Gegend der Inseln Maumere, Flores undKawe im Raja-Ampat-Archipel. Trimma xanthochrum n. sp.ist durch folgende Merkmale gekennzeichnet: breite Interor-bitalregion (80-100% des Pupillendurchmessers), einen zwei-ten Rückenstachel, der normalerweise nach hinten bis zwi-schen die Ansätze des zweiten und dritten Rückenflos-senstrahls reicht, 15-16 Brustflossenstrahlen, in der Regel mit7-8 verzweigten Strahlen, senkrechte Reihen von Sinnespa-pillen unterhalb vom Auge – mit jeweils 2-3 Papillen in denReihen 1 bis 4 und 4-5 Papillen in Reihe 5 –, einen Schwanz-fleck, dessen untere Hälfte etwa zwei Drittel der Breite deroberen Hälfte einnimmt, sowie normalerweise eine gelblicheGesamtfärbung auf dem Rumpf, die sich mindestens proxi-mal in einem Gelbton auf der Schwanzflosse fortsetzt.Gegenwärtig kennt man diese Art mit Sicherheit nur vomRaja-Ampat-Archipel.

RésuméUne récente collecte (2010) sur les Îles Raja Ampat d’Indo -

nésie a permis la capture de sept nouvelles espèces de Trimma,dont six sont décrites ici. Deux d’entre elles sont pour l’heureconnues exclusivement sur ces îles, tandis que les autres peu-vent être trouvées dans d’autres parties du Pacifique Ouest.Trimma cheni n. sp. se distingue des autres espèces de Trimmapar 8-9 écailles sur la ligne médiane prédorsale, 2-3 écaillessur l’opercule, un deuxième rayon épineux étiré sur la pre-mière nageoire dorsale allant à l’arrière jusqu’au-dessus desbases des rayons 1-4 de la seconde dorsale, des rayons de lanageoire pectorale divisés au milieu, le cinquième rayon de lapelvienne divisé une fois, une rayure sombre à la base desnageoires dorsales, l’attache des écailles indistinctement mar-quée d’une pigmentation plus sombre, et à l’état vivant, deuxbarres rouges à orange sur la joue et trois raies jaunes diffusessur le corps (les plus apparentes le long du pédoncule caudal).Elle a été recensée aux Palaos, aux Philippines, et au Sulawesiet à Flores en Indonésie. Trimma erdmanni n. sp. présente uncorps orange rougeâtre, avec une rayure latérale rougesombre, aux bordures plus foncées, qui remonte jusqu’à latête, où elle se divise ; là, une fine rayure longitudinale de cou-leur claire se trouve sous l’œil. Elle ne possède habituellementpas d’écailles sur la ligne médiane prédorsale, la seconde épinede la première dorsale est étirée, la dorsale compte 9 rayons etl’anale 8, une seule division sur le cinquième rayon de la pel-

aqua vol. 17 no. 3 - 10 July 2011 128

Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature

vienne et 19-22 branchiospines sur le premier arc branchial.L’espèce est connue sur les Îles Raja Ampat, les Îles Hermit età Madang (Papouasie-Nouvelle-Guinée), et sur des photogra-phies prises à El Nido (Île de Palawan) et dans le Golf deDavao (Mindanoro, Philippines). Trimma habrum n. sp. pos-sède un espacement interorbital osseux égal au diamètre de sapupille, 8-9 écailles sur la ligne médiane prédorsale, unenageoire pectorale à 14 rayons non divisés, une nageoire pel-vienne dont le cinquième rayon n’est pas divisé, une absencede membrane basale reliant le cinquième rayon de part etd’autre de la ligne médiane, habituellement une seule rangéecomplète de nageoires sur les joues et des écailles sur les deuxtiers supérieurs de l’opercule. Les tons frais de sa colorationsont diagnostiques : un dorsum pâle et translucide, avec destaches jaune clair et la base de chaque élément de la dorsaleentourée d’un point rouge, une barre rouge fine le long desbords postérieurs des hypuraux, une absence totale de pig-mentation foncée sur la région hypurale du pédoncule, et unepigmentation sombre couvrant les bordures dorsales de la ves-sie natatoire, la gaine des neurones et le cerveau. Elle est pourl’instant uniquement connue sur le résultat d’une seule col-lecte réalisée sur l’Île Kerou (Îles Fam, Raja Ampat, Indoné-sie). Trimma haimassum n. sp. se caractérise par un corps rela-tivement épais, de nombreuses écailles irrégulières avec envi-ron 30 rangées latérales et plus de 15 écailles dans les sériestransversales antérieures, un cinquième rayon de la pelviennese ramifiant deux fois de manière dichotomique, habituelle-ment une seconde épine dorsale quelque peu étirée, uneabsence de prédorsale, des écailles sur les joues ou les oper-cules, un espacement osseux interorbital limité avec uneexcroissance charnue médiane entre les deux yeux, et uneexcroissance dermique avant la première épine dorsale. Unpoint rouge ou une tache étirée au-dessus de l’opercule estvisible sur des spécimens vivants et frais, et la surface dorsaledu museau présente un motif réticulé foncé, avec des rayuressombres transversales sur la bordure dorsale de l’orbite. L’es-pèce est connue du nord du Sulawesi sud-occidental jusqu’àPalawan et l’est des Îles Salomon. Trimma papayum n. sp. estunique parmi les espèces décrites dans le genre pour avoir unpoint noir ocellé sur la pupille, d’un tiers du diamètre de celle-ci et un autre juste derrière le quatrième rayon épineux de ladorsale. Elle possède une nageoire dorsale à 9 rayons et uneanale à 8, un seul point de ramification sur le cinquièmerayon de la pelvienne, 10-11 rangées d’écailles transversalesantérieures et 8-9 postérieures, 5-10 écailles sur la lignemédiane prédorsale, et une seule rangée de 3 écailles sur labordure supérieure de l’opercule. Les spécimens fraîchementcollectés ont une coloration générale rouge orange, ponctuéede points jaunes diffus et éparpillés. L’espèce est seulementconnue en Indonésie, à Maumere (Flores) et sur l’Île Kawe(Raja Ampat). Trimma xanthochrum n. sp. est caractérisée parune large région interorbitale (80-100% du diamètre de lapupille), une deuxième épine dorsale arrivant habituellement,à l’arrière, entre les bases du deuxième et du troisième rayonde la dorsale, 15-16 rayons sur la pectorale dont habituelle-ment 7-8 sont divisés, des rangées verticales de papilles sensi-tives sous les yeux, de 2-3 papilles sur les rangées 1-4 et de 4-5 sur la rangée 5, une tache caudale dont la moitié inférieure

mesure deux-tiers de la moitié supérieure, et une colorationdu corps globalement jaunâtre, avec au minimum du jaune àproximité de la nageoire caudale. Pour l’instant, on ne l’aidentifiée avec certitude que sur les Îles Raja Ampat.

SommarioRecenti spedizioni nelle Isole Raja Ampat, Indonesia (2010)

hanno portato al campionamento di sette specie non descrittedi Trimma, sei delle quali sono descritte in questo articolo.Due di queste sono al momento note solo in queste isole, lealtre anche da altre parti del Pacifico occidentale. Trimmacheni n. sp. si distingue dalle altre specie di Trimma per avere8-9 scaglie sulla linea mediana predorsale, 2-3 scaglie sull’op-ercolo, la seconda spina dorsale allungata fino a raggiungereposteriormente la base dei raggi 1-4 della seconda pinna dor-sale, raggi pettorali mediani ramificati, quinto raggio pelvicocon una singola ramificazione, una stria basale scura sullepinne dorsali, avvallamenti delle scaglie indistintamentedelimitate da pigmento più scuro e, in vita, due barre rosso-arancio lungo le guance e sul corpo tre strie gialle diffuse (piùevidenti lungo il peduncolo caudale). È stata rinvenuta aPalau, Filippine e a Sulawesi e Flores in Indonesia. Trimmaerdmanni n. sp. ha il corpo arancio-rossastro con una stria lat-erale rosso scuro con bordi più scuri sul corpo che si esten-dono anteriormente sul capo, do ve si biforca, ed è presenteuna sottile striatura longitudinale chiara sotto l’occhio. Ingenere non possiede scaglie predor sa li, ha la seconda spinadella prima dorsale allungata, pos siede 9 raggi dorsali e 8anali, una singola ramificazione nel quinto raggio pelvico e19-22 rastrelli branchiali sul primo ar co branchiale. La specieè stata raccolta alle isole Raja Ampat e a Madang e alle isoleHermit (Papua Nuova Guinea) ed è stata fotografata a ElNido, isola di Palawan e nel golfo di Davao, Mindanoro, Fil-ippine. Trimma habrum n. sp. ha una distanza interorbitaleossea uguale al diametro della pupilla, 8-9 scaglie predorsali,14 raggi pettorali non ramificati, quinto raggio pelvico nonramificato, membrana basale che unisce il quinto pelvico allalinea mediana assente, in genere una singola fila completa discaglie sulla guancia e scaglie sui due-terzi superiori dell’oper-colo. La colorazione dell’esemplare fresco è diagnostica: ildorso è traslucente e pallido con macchie giallo chiaro e labase di ogni elemento della dorsale circondato da una mac-chia rossa, una sottile barra rossa lungo i margini posterioridegli ipurali, nessuna pigmentazione scura nella regione ipu-rale del peduncolo e una pigmentazione scura che ricopre imargini dorsali della vescica natatoria, la guaina neurale e ilcervello. Al momento è nota solo da un singolo campiona-mento eseguito a Kerou Island, Isole Fam, Raja Ampat,Indonesia. Trimma haimassum n. sp. si caratterizza per avereun corpo relativamente alto, numerose scaglie irregolari pre-senti in 30 file laterali più di 15 scaglie in linea anterioretrasversale, il quinto raggio pelvico che si ramifica due volte inmodo dicotomico, la se conda spina dorsale generalmenteallungata, assenza di scaglie predorsali e anche sulla guancia esull’opercolo, l’interorbitale moderatamente largo con unacresta mediana carnosa tra gli occhi e una cresta dermica ante-riore alla prima spina dorsale. È presente una macchia rossa ouna chiaz za allungata sopra l’opercolo negli esemplari appena

aqua vol. 17 no. 3 - 10 July 2011129



130aqua vol. 17 no. 3 - 10 July 2011

This paper describes six of those new species. Addi-tional specimens of some of these new species wereobtained from the Raja Ampat area by Mark Erd-mann on subsequent visits. The species describedhere were listed under their ‘RW’ numbers byDimara et al. (2010).

METHODSMethods and the format of the descriptions fol-

low Winterbottom (2002), except that pectoraland pelvic-fin ray branching is described from pre-served material stained with a cyanine blue solu-tion as outlined in Akihito et al. (1993; Akihito etal. 2002: 1270). Lengths given are StandardLength (SL) in millimetres; SD = Standard Devia-tion; values for the holotypes are given in boldwhere appropriate. In the additional (non-type)material examined sections, the catalogue numberis followed by the number of specimens in the lot(except where there is only a single specimen) withthe range of SL given in parentheses. Abbreviationsfor repositories of material examined follow thecodon abbreviations published by the AmericanSociety of Ichthyologists and Herpetologists(http://199.227.217.251/files/codons.pdf ). Theletters “CS” after a ROM catalogue number denotespecimens that were cleared with trypsin andcounter-stained with alizarin and alcian blue. Thevertebral transition type (A or B) between theabdominal and caudal vertebral centra is definedand illustrated by Winterbottom (2003: Fig. 14(inset)), and Winterbottom & Zur (2007: Fig. 3).Essentially, in type A, the last two abdominal ver-tebrae are of the usual perciform configuration inthat they lack a bony connection across the midlinebetween the bases of the paired haemal arches, andthe first (and often subsequent two to three) caudalvertebrae have a single greatly enlarged canal withonly the tips of the haemal arches fused together inthe midline. In type B, the last two abdominal ver-tebrae have a bridge joining the haemal archesacross the midline, forming a haemal canal; thefirst caudal vertebra has broad haemal arches witha small basal haemal canal and a larger secondarycanal distal to this, which forms a posteriorly taper-ing funnel and embraces the posterior end of theswimbladder. Counts and measurements wereinput directly into an Excel file with Mitutoyo dig-ital callipers using WinWedge 3.01™ software.Photographs other than the portraits of fresh orlive specimens were produced from multiple digitalimages taken with a Nikon D100 or D300S cam-

raccolti, mentre la superficie dorsale del muso mostra unmotivo reticolato scuro, con striature trasversali scure sopra ilmargine dorsale dell’orbita. La specie è nota dal Sulawesi sud-occidentale a nord fino a Palawan e a est fino alle IsoleSalomone. Trimma papayum n. sp. è unica tra le specie de -scritte in questo genere per avere una macchia ocellata neradal diametro pari a un terzo a quello della pupilla sopra e ap -pena dietro la quarta spina dorsale. Possiede 9 raggi dorsali e8 anali, un singolo punto di ramificazione nel quinto raggiopelvico, 10-11 file di scaglie trasversali anteriori e 8-9 po -steriori, 5-10 scaglie predorsali e una sola fila di 3 scaglie sulmar gine superiore dell’opercolo. Esemplari appena raccoltihanno una livrea nel complesso rosso-arancio, con mac chiegialle diffusamente disseminate. La specie è nota solo inIndonesia, a Maumere, Flores e Kawe Island, Raja Ampat.Trimma xanthochrum n. sp. è caratterizzata da un’ampiaregione interorbitale (80-100% del diametro della pupilla),da una seconda spina dorsale che raggiunge di solito posteri-ormente lo spazio tra le basi del secondo e il terzo raggio dor-sale, 15-16 raggi pettorali con generalmente 7-8 raggi ramifi-cati, file verticali di papille sensoriali sotto l’occhio formate da2-3 papille nelle file 1-4 e 4-5 nella fila 5, una chiazza caudaleche ha la metà inferiore circa due terzi l’ampiezza della metàsuperiore e, infine, una colorazione generalmente giallastracon del giallo almeno prossimalmente nella pinna caudale.Attualmente è nota con certezza solo alle Isole Raja Ampat.

INTRODUCTIONTrimma Jordan & Seale (type species: T. caesiura

Jordan & Seale, 1906) contains about 100 species ofsmall (<30 mm SL), often colourful gobiids, primar-ily associated with Indo-Pacific coral reefs. Membersof the genus may be recognized by the lack ofcephalic sensory canal pores, a much reducedcephalic sensory papillae pattern, a wide gill openingextending anteriorly to below the vertical limb of thepreopercle or, more usually, anterior to this, a lack ofspicules (odontoids) on the outer gill rakers of thefirst gill arch, fewer than 12 dorsal and anal fin rays,and a fifth pelvic-fin ray that is equal to or more than40% the length of the fourth pelvic-fin ray (Winter-bottom 1995, 2003). Although the gill rakers on thefirst gill arch of members of this genus often have arow of crenulations or serrations (a series of sharplyrounded protrusions lacking any ossification) alongtheir medial margins (Fig. 1), these structures are notconsidered here as the equivalent of odontoids. Thereare currently 64 valid species of Trimma, withapproximately 35 additional known undescribedspecies (Winterbottom & Hoese, unpublished).A recent (January/February, 2010) rapid survey of

the Raja Ampat islands off the Bird’s Head region ofNew Guinea sponsored by Conservation Interna-tional’s Indonesia Marine Program resulted in thecollection of seven undescribed species of this genus.

Table I. Counts of cephalic sensory papillae in the different rows of the six new species of Trimma. Values consist of mini-mum, maximum, with the mean, Standard Deviation and number of specimens examined respectively given in parentheses,except where the value was invariate, where only the number of specimens is given.

Papillae row T. cheni T. erdmanni T. habrum T. haimassum T. papayum T. xanthochrum

Row a 4-5 (4.9, 0.21, 20) 5-6 (5.1, 0.22, 19) 2-3 (2.7, 0.45, 11) 5 (20) 5 (9) 4-6 (4.7, 0.56, 18)

Row b 4-7 (5.8, 0.67, 17) 5-9 (7.7, 1.03, 19) 3-6 (4.8, 0.83, 8) 9-13 (10.9, 1.03, 16) 6-11 (7.6, 1.28, 10) 4-7 (5.6, 0.8, 10)

Row c 5 (20) 5-6 (5.1, 0.22, 19) 4-5 (4.8, 0.39, 11) 6 (20) 5-6 (5.1, 0.31, 9) 5-6 (5.9, 0.22, 19)

Row d 6-8 (6.8, 0.55, 19) 7-12 (9.1, 1.43, 19) 4-6 (5, 0.63, 10) 11-16 (12.8, 1.26, 20) 7-10 (8.5, 1.02, 10) 6-8 (6.7, 0.75, 18)

Row d’ 7-10 (7.9, 0.92, 20) 8-12 (10.1, 1.02, 19) 5-7 (5.3, 0.62, 11) 10-16 (11.9, 1.48, 20) 9-11 (9.9, 0.7, 10) 6-10 (7.6, 1.07, 20)

Row e –ant. 11-18 (13.5, 1.63, 19) 13-20 (16.5, 2.11, 19) 11-15 (11.8, 1.28, 12) 16-23 (19.1, 1.89, 20) 15-19 (16.2, 1.08, 10) 12-19(14.5, 1.67, 19)

Row e –pos. 11-17 (14.1, 1.71, 19) 15-23 (19.2, 1.85, 19) 12-16 (13.3, 1.48, 11) 15-26 (20.7, 2.31, 20) 15-18 (16.9, 1.04, 10) 9-23 (17.2, 2.79, 17)

Row i – ant. 7-9 (7.7, 0.52, 19) 6-9 (7.3, 0.71, 19) 6-7 (6.6, 0.48, 11) 8-11 (9.3, 0.71, 20) 7-8 (7.6, 0.5, 9) 6-7 (6.9, 0.31, 19)

Row i – pos. 7-9 (8, 0.44, 20) 8-10 (8.2, 0.52, 19) 4-8 (6.5, 1.08, 11) 9-11 (10, 0.67, 20) 7-8 (7.4, 0.5, 9) 7-8 (7.7, 0.44, 19)

Row p 6-7 (6.9, 0.21, 20) 6-7 (6.1, 0.31, 19) 6-9 (8.2, 0.94, 11) 8 (20) 6-7 (6.4, 0.5, 9) 8-9 (8.3, 0.46, 20)

Row r 2 (20) 2 (19) 2 (11) 2 (20) 2 (9) 4-5 (4.6, 0.49, 20)

Row f 2-3 (2.9, 0.21, 20) 3-4 (3.1, 0.22, 19) 2-3 (2.9, 0.3, 10) 3-5 (3.9, 0.44, 20) 3-4 (3.4, 0.48, 11) 3-6 (4.1, 0.94, 19)

Row cs” 2-3 (2.9, 0.21, 20) 3 (19) 2-3 (2.9, 0.3, 10) 3-4 (4, 0.22, 20) 3 (9) 2-5 (3.4, 0.91, 20)

Row g Apparently absent 4-10 (8.1, 1.44, 15) Apparently absent 11-17 (12.2, 1.61, 14) 5-9 (7, 1.6, 7) Apparently absent

Row x 6-9 (6.8, 0.67, 20) 7-10 (8, 0.79, 19) 6-9 (7.8, 0.92, 9) 8-11 (8.7, 0.84, 20) 6-8 (7.4, 0.68, 9) 6-9 (7.9, 0.86, 16)

Row z 5-8 (6.4, 0.8, 10) 6-11 (8.1, 1.35, 18) 6-7 (6.4, 0.48, 8) 7-9, (7.9, 0.65, 20) 7-10 (7.6, 0.96, 9) 5-7 (5.8, 0.83, 4)

Row ot 11-16 (13.8, 1.6, 10) 15-20 (17.6, 1.38, 18) 12-15 (12.8, 1.03, 9) 16-20 (18.1, 1.24, 20) 14-19 (16.1, 1.22, 10) 13-19 (15.9, 1.54, 13)

Row os 4-6 (4.7, 0.62, 9) 6-9 (7.9, 1.09, 15) 5-7 (6, 0.63, 5) 6-9 (7.5, 0.92, 20) 7-10 (8, 1.05, 9) 6 (1)

Row oi 2-4 (3.2, 0.6, 10) 4-7 (5.2, 0.71, 17) 3-4 (3.1, 0.33, 8) 4-7 (5.3, 0.92, 19) 2-5 (4.2, 0.87, 10) 4-7 (5.2, 1.05, 15)


131 aqua vol. 17 no. 3 - 10 July 2011

lows the orientation of, the maxilla and for a hori-zontal row across the lower cheek that is, at least inTrimma, well separated from the first row anteri-orly. Miller (1972a) refined this nomenclature,using d’ for the row behind the maxilla, and reserv-ing d for the row across the cheek, and this dis-tinction is followed in this paper. [Note: Miller(1972a; 1972b) actually uses d and d1 respectivelyin the text of both papers and for the figure in the1972b paper, but employs d’ in Figure 5 of the1972a publication. Since Sanzo (1911) consis-tently uses “prime” where appropriate for otherrows of papillae, I have opted to follow the nomen-clature used in Miller’s figure (1972a, Fig. 5)]. Thisseparation of row d into two discrete entities is alsofound in both Priolepis and Egglestonichthys (Miller

era attached to a Zeiss SV-8 dissecting microscopeat slightly incremental focal planes selected manu-ally, or with a Canon EOS Rebel XS cameraattached to a Zeiss SV-12 dissecting microscopeusing Zeiss AxioVision 4.8™ software and auto-matic increments. The images were then collatedinto a single image using Helicon Focus 5.1™(HeliconSoft). All images, drawings and image col-lations were made by the author unless otherwisenoted. Type specimens are confined to those col-lected in the Raja Ampat islands of Indonesia.Cephalic sensory papillae: Nomenclature for the

cephalic sensory papillae follows Sanzo (1911),with the exception of his row d on the lateral sur-face of the cheek. Sanzo (1911) uses this letter forboth the row immediately behind, and which fol-

Fig. 1A-C. Left lateral view of the upper portions of thefirst gill arches to show crenulated edges of gill rakers in: A.Trimma erdmanni (26.2 mm SL paratype, ROM 85033 –image reversed); B. T. haimassum (28.8 mm SL paratype,ROM 85347); and C. T. haimassum (27.3 SL paratype,ROM 1836CS). A and B stained with cyanine blue, Ccleared and stained with alizarin and alcian blue – notelack of crenulations, indicating that they are structureswithout any calcification.

Fig. 2A-B. Composite image of head to show distributionand nomenclature of papillae. A. Trimma haimassum(upper = dorsal view of left half, lower = left lateral view);B. T. xanthochrum (left lateral view of cheek area). The greyconnecting lines in A represent the rows as counted in thispaper, those in B represent parts of the nerves that stainedin the cyanine blue preparation.


132aqua vol. 17 no. 3 - 10 July 2011

the anterior and posterior sections respectively).However, such a separation is apparently rare ingobies (D. F. Hoese, in litt.). The complete nomen-clature of the papillae rows used here is given inFigure 2A. Sanzo’s (1911) nomenclature waslargely accepted and used by Aurich (1938),although a brief comparison of the two worksbrings several apparent discrepancies to light.Hoese (1983) pointed out some of the difficultiesin homologizing Sanzo’s (1911) names with thepapillae patterns he found in Indo-Pacific gobies,and proposed a revised nomenclature based onwhether the bases of the papillae were longitudi-nally or vertically oriented with respect to the axisof the papilla row. Takagi (1988) also proposed a

& Wongrat 1979; Winterbottom & Burridge1992). It is also present in the possibly relatedLythrypnus (Ahnelt & Bohacek (2004) - where theauthors referred to the two sections as d1 and d2 for


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lae. It seems probable that when homologies can beempirically established, a single, unified (and differ-ent) nomenclature for papillae will need to beadopted. The recent explorations of the innervationsof the different papillae patterns in various gobioids(e.g. Ahnelt & Bohacek 2004; Asaoka et al. 2011) isa promising approach, but also raises some of the dif-ficulties envisioned by Hoese (1983). For example,the phylogenetic homology of the papillae in therows a, c and cp may prove especially difficult toresolve. Most of the head papillae have an ovoid base,and the long axis is nearly always oriented at rightangles to the path of the nerve supplying them.Apparent exceptions to this in most species ofTrimma, such as rows a, c and p, where the path ofthe nerve is aligned with the long axis of the bases ofthe papillae, may well prove to be the result of reduc-tion to a single papilla of originally transverse rowsthat had the papillae bases transversely oriented withrespect to their immediate nerve track. Since the papillae patterns in Trimma correspond

well, for the most part, with those illustrated andlabelled by Sanzo (1911: Pl. 9, Figs 9-10) for Gobiusaffinis Kolombatovic, 1891 (current status:Pomatoschistus pictus adriaticus Miller, 1973), Sanzo’snomenclature is employed here with the exceptionnoted above (Fig. 2A). Although the identity of allthe rows of papillae labelled in figures of Trimma inthis paper appear to be positional homologues ofSanzo’s rows, they should not be construed as phylo-genetic homologues. Note also that the grey linelinking individual papillae into the named rows asused in this paper (Fig. 2A) is not based on an exam-ination of the innervations or of the phylogenetichistory of the genus, and it is probable that somepapillae are incorrectly attributed to or omitted fromthat row. The papillae are easily abraded from thesurface of the head (especially those in rows b, d, zand g), and are usually difficult to see where the areais covered with scales. Where counts are given, theyshould thus be regarded as minimum values. Thosecounts exhibiting large variation probably do sobecause some of the papillae have been lost.

Trimma cheni n. sp.(Figs 3-6)Face-stripe pygmy goby

Trimma RW sp. 51 – Dimara et al. 2010: 621(Raja Ampat)

Trimma sp. 7 – Allen & Erdmann 2009: 619(western Bird’s Head Peninsula)

system of nomenclature for the papillae rows, andprovided a table equating his names with thoseused by previous authors (Takagi 1988: Tbl. 4).Other authors have been content to simply illus-trate the papillae patterns (although Akihito et al.(1988: Fig. 36) did distinguish between the trans-verse and longitudinal rows). Trimma has beencharacterized as having a reduced transverse papil-lae pattern, consisting only of longitudinal rows ofpapillae on the cheek (Winterbottom & Burridge1992; Hoese et al. 2011). One of the putative sis-ter groups of Trimma, Priolepis (Winterbottom1984; Miya, pers.comm.) has several transverserows of cheek and interorbital papillae, at least inthe hypothesized basal members of that clade(Winterbottom & Burridge 1992: Fig. 12). Mostspecies of Trimma have two longitudinal rows ofpapillae (a and c) on the cheek immediately underthe eye and above row d. Frequently, the anterior-most papillae in rows a and c lie below the anteriorborder of the eye forming a vertical row of two papil-lae, and the papillae below the posterior margin ofthe pupil form a vertical row of two or three papillae(in the later case potentially attributable, from dorsalto ventral, to one papilla from each of rows a and cand a single papillae named cp by Sanzo (1911),respectively). Ahnelt & Bohacek (2004) also employthis nomenclature for the ventralmost papilla inLythrypnus, but Asaoka et al. (2011: Fig. 4) use cp forthe entire vertical row, which consists of at least five,perhaps more, individual papillae in Pterogobiuselapoides. Ahnelt & Göschel (2003: Fig. 3) also use cpfor a vertical row of several papillae on the cheek inQuietula. One of the new species described here(Trimma xanthochrum) has two to five papillae ineach of five vertical rows in this position. Where suchvertical rows occur, Sanzo (1911) employs numbers1 through 6 as identifiers, beginning with “1” for therow below the anteroventral margin of the eye. Thefifth row, below the posterior margin of the pupil,usually marks the anterior limit of row b. Given thehypothesis that Trimma exhibits a reduced transversepattern of cheek papillae (with T. xanthochrum pos-sessing the most plesiomorphic condition in thegenus), the application of Sanzo’s (1911) nomencla-ture of rows a and c for the longitudinal papillae rowsbelow the eye is incorrect since these papillae repre-sent the remnants of the vertical rows 1-5 (see alsoHoese et al. 2011). An alternative nomenclature isgiven in Figure 2B, based on an interpretation thatthe papillae rows below the eye in Trimma do repre-sent originally more extensive vertical rows of papil-

Fig. 3. Left lateral view of Trimma cheni, 22.3 mm SL male paratype, ROM 85085, Mutus Island, Raja Ampat. Photo by R.Winterbottom.

Fig. 4. Left lateral view of Trimma cheni (live), El Nido, Palawan, Philippines. Photograph by G. R. Allen.

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2010, M. V. Erdmann. MZB 19776 (formerlyROM 87479), 10, 11.1-21.9, Waigeo Island, Tan-jung Manare (00°15’26.1”S, 130°19’01.5”E), 52 m,clove oil, 28 May, 2010, M. V. Erdmann. ROM85085, 4, 14.3-22.3, about 1.2 km SSE of MutusIsland, on west side of sand spit (00°21’ 01.0”S,130°21’25.4”E), 50 m, clove oil, 0900-0920, field# RW10-16, 28 January 2010, M. V. Erdmann.ROM 85138, 11, 10.2-19.2, Jef Tsiep Island, westside, channel between it and small island to thewest (00°23’05.7”S, 130°16’37.1”E), 19.8-22.9m, clove oil, 1600-1630, field # RW10-20, 28 January 2010, R. Winterbottom, L. Katz & P.Johannes. ROM 85161, 17, 10.9-21.8, TanjungManare, Waigeo Island, west side at about middle of width off a small cape (00°16’26.1”S,130°19’01.5”E), 19.8-22.9 m, rotenone, 0750-0820, field # RW10-23, 29 January 2010, R. Win-

Material Examined: A total of 8 lots, 56 typespecimens (10.2-22.3 mm SL), plus an additionalnon-type specimen (tissue voucher), all from theRaja Ampat Islands, Indonesia, and other speci-mens from Indonesia and the Philippines. Thedescription is based on the holotype and 19 otherspecimens from ROM 85161, 85138, 85179 and1835CS, 16.2-22.1 mm SL (x = 19.3, SD = 1.31),12 males and 8 females.Holotype: ROM 87481, 20.0 mm SL male, Tan-

jung Manare, Waigeo Island, west side at aboutmiddle of width off small cape (00°16’26.1”S,130°19’01.5”E), 52 m, rotenone, 0750-0810, field# RW10-24, 29 January 2010, M. V. Erdmann.Paratypes: AMS I.45590-001 (formerly ROM

85221), 3, 19.5-21.5, Wofoh Island, west coast nearsouth end (00°15’21.9”S, 130°17’32.0”E), 48 m,clove oil, 1610-1650, field # RW10-26, 29 January

Fig. 5. Left lateral view of Trimma cheni, 17.6 mm SL juvenile paratype, ROM 85315, Waaf Island, Raja Ampat. Note theparasitic copepod attached immediately posterior to pelvic fins. Photo by R. Winterbottom.

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teriorly to bases of rays 1-3-4 of second dorsal finwhen adpressed (x = 2.3, SD = 1.02, n = 19), raysall branched except, usually, posterior element oflast ray (first ray unbranched in 2), last ray reach-ing posteriorly 40-50% of distance from its base tofirst dorsal procurrent ray (x = 48.8, SD = 3.05);anal fin I 8, all but first and posterior element ofposteriormost ray branched; posteriormost ray 35-40-45% of distance from its base to first ventralprocurrent ray (x = 41.0, SD = 2.55, n = 19); pec-toral fin 16-17-18 (x = 17.1, SD = 0.38), 4-5-7(mean = 5.4, SD = 1.03) and 4-8 (x = 5.7, SD =1.36) unbranched dorsal and ventral rays respec-tively, middle rays branched, fin reaching posteri-orly to a vertical line between bases of anal rays 1-3 (x = 1.3, SD = 0.56, n = 19); pelvic fin I 5, nofrenum, basal membrane reduced and 10-15 %length of fifth ray (x = 13.0, SD = 1.91), first fourrays with one sequential branch, fifth ray branchedonce and 48-52-56% length of fourth (x = 52.6,SD = 2.50, n = 18), fourth ray reaching posteriorlyto between bases of anal rays 1-3.5-6 (x = 4.0, SD= 1.24, n = 17). Lateral scales 23; anterior trans-verse scales 7-8-8.5 (x = 8.1, SD = 0.37); posteriortransverse scales 7-8-8.5 (x = 7.5, SD = 0.41); pre-dorsal scales 8-9 (x = 8.2, SD = 0.39); no scales oncheek; opercle with single row of 2-3 usuallyctenoid scales; pectoral base with usually 3 verticalrows of scales and 4 cycloid scales in posterior row;5-7-8 (x = 6.4, SD = 0.79) cycloid prepelvic scales(in midline anterior to basal membrane); 11-12 (x= 11.9, SD = 0.3) circumpeduncular scales; bodyscales ctenoid except for cycloid scales on anteriorbelly midline and on, beneath and just posterior topectoral fin base; body scales extending anteriorlyto just behind eye. Gill opening extendinganteroventrally to a vertical below anterior one-third of pupil. Upper jaw with outer row of curved,enlarged, spaced canines along length of premax-

terbottom, R, L. Katz, P. Johannes, W. Kaka & W. Awom. ROM 85197, 7, 13.4-19.8, collectedwith the holotype. ROM 85315, 1, 17.6, S of Misool Island, off Waaf Island (02°08’56.4”S,130°13’17.0”E), clove oil, 0715-0740, field #RW10-36, 1 February 2010, M. V. Erdmann.ROM 1835CS (formerly ROM 85232), 2, 19.6-21.2, Wofoh Island, west coast near south end(00°15’21.9”S, 130°17’32.0”E), 12-16 m, cloveoil, 1610-1700, field # RW10-27, 29 January2010, L. Katz. Additional (Non-type) Material. Raja Ampat:

ROM T07737, (21.8), collected with ROM 85085(tissue specimen). Indonesia: Flores, ROM 62652,2 (17-19). Sulawesi, ROM 64641, 2 (19-21).Palau: Ulong Pass, ROM 74805, 5 (10-18).Philippines: Cebu, ROM 49247, 8(9-19); ROM53114, 3 (20-21). Negros Oriental, ROM 53112,9 (12-22); ROM 53113, (18). Siquijor, ROM53107, (19); ROM 53108, 6 (19-22); ROM53109, 8 (16-21); ROM 53110, 11 (10-22); ROM53111, 8 (21-24).

Diagnosis: Trimma cheni has 8-9 scales in thepredorsal midline, a row of 2-3 usually ctenoidscales along the upper border of the opercle, nocheek scales, a dark basal stripe in the dorsal fins,an elongate second dorsal spine reaching posteri-orly to the bases of rays 1-4 of the second dorsalfin, the middle rays of the pectoral fin branched, afifth pelvic fin ray which branches once dichoto-mously, the basal membrane connecting the innermargins of the fifth pelvic fins rays is less than 15%the length of the fifth ray, the scale pockets are notdistinctly outlined with darker pigment, and, inlife, there are two red to orange bars across thecheek and three diffuse yellow stripes on the body(most obvious along the caudal peduncle).

Description: Dorsal fins VI + I 8-9 (x = 9.0, SD= 0.22), second third spine longest, reaching pos-

Fig. 6. Map showing the distributions of the new species of Trimma described. ☒ all species present; △ – T. cheni; ◘ T. erd-manni; ✪ T. haimassum; ♦ T. papayum; ★ T. xanthochrum.

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Bony interorbital 31-35-44% (x = 38.1, SD = 4.08)pupil width, with broad moderately developed U-shaped interorbital furrow and shallow postorbitalgroove (or none). No ridge of tissue (or dermalcrest) extending anterior to first dorsal spine. Epax-ialis musculature extending anteriorly to above ver-tical with posterior margin of pupil. Last twoabdominal vertebrae with bridge of bone connect-ing haemal arches just distal to their bases, forming‘haemal canal’ bounded dorsally by centrum. Firstcaudal vertebra with two canals, a small proximalone at base of haemal arches, which fuse in midlinebriefly before diverging again to form posteriorlydirected funnel-like second arch. Haemal archesthen fusing again in midline to form haemal spine(vertebral transition Type B).

C o l o u r p a t t e r n (from images of four freshlycollected specimens, 17.6-22.3 mm SL). A 22.3mm SL male (Fig. 3) has a mostly red head, darkeron the snout, with two lighter and more intensered bars with irregular diffuse yellowish margins onthe cheek below the anterior and middle of the eye,and another yellow-brown bar over the verticallimb of the preopercle, the dorsal half of the oper-cle with a similar colouration; the posterior margin

illa, gradually decreasing in size posteriorly untilnot much larger than inner rows; two to three irreg-ular inner rows of smaller teeth almost reaching dis-tal tip of premaxilla, innermost row of slender,slightly curved, spaced teeth almost as long as outerrow, from symphysis to bend of jaw in males andsome females. Lower jaw with outer row of 4-5curved, enlarged, spaced canines reaching to bendof dentary, several irregular inner rows of slightlycurved caniform teeth half size of outer, becomingreduced in size and less curved posteriorly andtapering to form single row of straight conical teethat beginning of coronoid process; innermost row ofenlarged slightly curved teeth at symphysis, increas-ing in size along jaw (equal in size to outermost rowjust behind bend of dentary, then gradually decreas-ing in size to dorsal tip of coronoid process (becom-ing equal in size to teeth in inner rows). Cephalicsensory papillae row counts given in Table I.Tongue truncate with rounded edges. Gill rakers onfirst arch 3-4 + 15-16 = 18-19-20 (x = 19.2, SD =0.59). Anterior nasal opening a short tube extend-ing out over upper lip, posterior nasal opening apore with raised rim, both protruding from slightlyraised oval sac confined to anterior half of snout.


137 aqua vol. 17 no. 3 - 10 July 2011

ent. The dark basal stripes in the dorsal and analfins remain obvious, the dark stripe at the mid-region of the second dorsal fin is relatively welldefined, and three diffuse dark stripes run parallelto the rays of the caudal fin. The dorsal rim of thepectoral fin base is often somewhat darker than thesurroundings, appearing as a diffuse dark streak.Comparisons: Trimma cheni belongs to a group of

species which has a scaled predorsal midline, nocheek scales, a scaled opercle, at least the middlerays of the pectoral fin are branched, and a darkbasal stripe in the dorsal fins is present (Winterbot-tom 2006). Within this group, T. cheni appearsclosest morphologically to T. milta Winterbottom,2002, in having a poorly developed postorbitaltrough or none and in possessing a single row of 2-3 scales on the opercle. However, T. milta does nothave the second dorsal spine elongated, has anunbranched fifth pelvic fin ray, has the scale pock-ets on the body lightly outlined with darker pig-ment, and lacks the conspicuous red to orange barson the cheek as well as the three yellow stripes onthe body. Trimma anthrenum Winterbottom, 2006,differs in lacking opercular scales and elongate sec-ond dorsal spine, in possessing a well developedpostorbital trough, and in live colouration (plainyellow with a distinctively coloured iris (see Win-terbottom 2006: Figs 1-2). Trimma preclarum Win-terbottom 2006 differs from T. cheni in lackingscales on the opercle, in possessing a well developedpostorbital trough, and in lacking the two red toorange bars on the cheek, although this species doeshave three yellow stripes on the posterior part of thebody. These stripes are also evident in T. hayashiiHagiwara & Winterbottom, 2007, which lacksscales in the predorsal midline (or, if scales are pre-sent, they are represented one or two scales crossingthe midline between the eye and the first dorsal finorigin) and on the opercle, and has a distinctiveblack ocellated spot on the branchiostegal mem-branes below the vertical limb of the preopercle.Distribution: Trimma cheni is currently known

from the Raja Ampat islands, and from Palau, thePhilippines and south-west to Sulawesi and Floresin Indonesia (Fig. 6) at depths of 5-52 m.Etymology: The species is named for I.-S. Chen,

National Taiwan Ocean University, Keelung, Tai-wan, in acknowledgment of his numerous publica-tions on Indo-Pacific reef fishes in general, andgobies in particular. This species has been referredto as Trimma RW sp. 51.

of the branchiostegal membrane immediately adja-cent to the opercle and subopercle is orange. Theiris is red grading to yellow orange ventrally, withan anteroventral to posterodorsal dark purple stripeacross the pupil; the dorsal surface of the orbit hasseveral alternating red and white stripes passingacross the midline. The body is yellowish, gradingto pink on the peduncle, with three vague yellowstripes (most obvious on the peduncle), with heav-ily scattered dark chromatophores, especially onthe dorsum and along the scale pockets and scalemargins. The membranes of the dorsal fins aredensely speckled with melanophores, except for ahalf-pupil diameter wide yellow stripe just abovethe base and, above this, a second but more diffuseyellow stripe. The membranes of the caudal fin aresimilarly invested with melanophores, with severalstreaks and blotches of yellow separated by threedarker diffuse stripes. The distal margin of the analfin is dusky, and there is a broad (about one-and-a-half pupil diameters wide) yellow stripe with dif-fuse margins and a dark basal stripe. The pectoraland pelvic fin rays are yellow or orange, with hya-line membranes, the bases of the fins are light tanwith many scattered dark brown chromatophores.A 20.1 mm SL male is similar, but with a darkerand greyer body, while a 20.0 mm SL male is paler,with a generally more yellow cast. A specimen fromEl Nido, Philippines photographed underwater(Fig. 4) has a greyish background with diffuse yel-low-green markings on the body, three bars on thelower part of the head (two below the eye, the thirdover the vertical limb of the preopercle), whichgrade from orange yellow dorsally into the green-ish-yellow ventrally in the region of the lower jawand branchiostegals. There are two short obliqueorange bars passing posterodorsally from the poste-rior margin of the upper orbit, and three shortorange to red transverse bars across the dorsal mar-gin of the eye. Other specimens photographedunderwater have a redder head and either a green-ish-yellow or brownish body. A 17.6 mm SL juve-nile (Fig. 5) has a relatively dark body, two dark redbars on a grey-pink cheek, and the stripes in thefins are reddish rather than yellow.

C o l o u r p a t t e r n i n a l c o h o l : straw-coloured with a heavy scattering of brown chro-matophores and melanophores on the body, napeand pectoral fin base. The opercle has numerousdensely scattered melanophores. The cheek is paler,and two diffuse lighter bars (corresponding to thered bars in fresh material) may or may not be appar-

Fig. 7A-B. Head of Trimma erdmanni (19.2 mm SL femaleparatype, ROM 85033) in: A. left lateral and B. dorsalview. Specimen stained with cyanine blue. Photo by R.Winterbottom.


138aqua vol. 17 no. 3 - 10 July 2011

Kawe Island, SW bay (00°05’15.2”S, 130°07’25.3”E), 50 m, clove oil. M. V. Erdmann, 26 Jan-uary 2010.Paratypes: AMS I.45591-001 (formerly ROM

85330), 1, 23.1, Kepotsol Island, east side(02°09’32.1” S, 130°17’34.0” E), 66 m, clove oil,field # RW10-38, 0830-0850, 1 February 2010,M. V. Erdmann. MZB 19777 (formerly ROM87460), 7, 11.2-22.5, Waigeo Island, TanjungManare (00°15’26.1”S, 130°19’01.5”E), 52 m,clove oil, no field No., 28 May, 2010, M. V. Erd-mann. ROM 85033, 22, 10.2-26.2, collected withthe holotype. ROM 85144, 8, 12.3-23.5, Jef TsiepIsland, west side, channel between it and smallisland to the west (00°23’05.7”S, 130°16’37.1”E),42 m, clove oil, field # RW10-21, 1600-1620, 28January 2010, M. V. Erdmann. ROM 85186, 16,10.5-22.3, Tanjung Manare, Waigeo Island, westside at about middle of width off small cape,sponges, hard corals (00°16’26.1”S, 130°19’01.5”E), 52 m, 1 kg powdered rotenone & detergent,field # RW10-24, 0750-0810, 29 January 2010,M. V. Erdmann. ROM 85393, 7, 15.2-22.2, SEislands off Misool, south side of Balbulol Island(02°01’29.5”S, 130°41’34.9”E), 45 m, 1 kg pow-dered rotenone & detergent, field # RW10-45,0750-0810, 2 February 2010, M. V. Erdmann.ROM 1837CS, 4, 18.2-23.0, Wofoh Island, westcoast near south end (00°15’21.9”S, 130°17’32.0”E), 48 m, clove oil, field # RW10-26, 1610-1650,29 January 2010, M. V. Erdmann.

Additional (Non-type) Material. Raja Ampat:ROM T07723 and T07724, 2 (24.1-24.1), col-lected with ROM 85033; ROM T07743 andT07744, 2 (7.0-21.1), collected with ROM85144. Papua New Guinea. Hermit Islands:USNM 243952, 17 (13-21. Madang: WAMP.30358-008, (19.0); WAM P. 30369-011, (21.2).Diagnosis: Trimma erdmanni differs most tren-

chantly from other species of the genus in thecolour pattern of a red to orange body with adarker red to orange lateral stripe with dark borderson the body which extends anteriorly onto thehead, where it bifurcates, and there is a thin lightstripe from the maxilla to the opercle that abuts theventral margin of the eye. The new species usuallylacks scales in the predorsal midline, but whenthese are present, they are separated from the firstdorsal fin by an unscaled area. The second (andthird to some degree) spines of the first dorsal areelongated, there are 9 dorsal and 8 anal rays, themiddle rays of the pectoral fin are branched, there

Trimma erdmanni n. sp.(Figs. 1A, 6-12)Erdmann’s pygmy goby

Trimma anaima Winterbottom 2000 – Allen &Erdmann 2009: 619 (in part, Bird’s Head Penin-sula)


Trimma sp. – Allen et al. 2003: 329 (lower right),Philippines

Material Examined: A total of 7 lots, 64 typespecimens, plus four additional non-type speci-mens (tissue vouchers), and three additional lots ofnon-type specimens from Papua New Guinea. Alltype specimens collected at the Raja AmpatIslands, Indonesia. Description based on up to 24specimens from ROM 85033, 85144, 85186,87460, 87484 and 1837CS (18.2-26.2 mm SL, 12males, 12 females). Holotype: ROM 87482, 23.5 mm SL male,

Fig. 8. Left lateral view of Trimma erdmanni, 26.2 mm SL male paratype, ROM 85033, Kawe Island, Raja Ampat. Note thatthe spine and first ray of the second dorsal fin are slightly deformed. Photo by R. Winterbottom.

Fig. 9. Left lateral view of Trimma erdmanni (live), Kawe Island, Raja Ampat. Photo by M. V. Erdmann.

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of distance between its base and first dorsal procur-rent caudal fin ray; anal fin I 8 (n = 24), all but firstand posterior element of posteriormost raybranched, last ray extending between 40-80% (x =60.7, SD = 12.4) of distance between its base andfirst ventral procurrent caudal fin ray; pectoral fin17-18-19 (x = 18.0, SD = 0.61), 3-4 (x = 3.4, SD= 0.49) and 2-4-7 (x = 3.4, SD = 1.00) unbrancheddorsal and ventral rays respectively, middle raysbranched, fin reaching posteriorly to a vertical linebetween anterior margin of urogenital papilla andbase of first ray of anal fin; pelvic fin I 5, nofrenum, basal membrane poorly developed, 7-17-

is a single branch in the fifth pelvic fin ray, thebasal membrane is 7-18% of the length of the fifthfin ray and there are 19-22 gill rakers on the outermargin of the first gill arch.Description: Dorsal fins VI + I 9, second and

third spines longest, second spine reaching tobetween bases of second to eighth rays of seconddorsal fin, third spine reaching to between bases ofdorsal spine and fourth ray of second dorsal finwhen adpressed, first ray of second dorsal fin usu-ally unbranched (branched in 5, SD = 0.41), pos-terior element of last ray unbranched, last rayextending between 50-100% (x = 74.2, SD = 16.0)

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Fig. 10. Left lateral view of Trimma erdmanni, 23.5 mm SL male holotype, ROM 87482, Kawe Island, Raja Ampat. Photoby R. Winterbottom.

Fig. 11. Trimma erdmanni, 14.3 mm SL juvenile paratype, ROM 85033, Kawe Island, Raja Ampat. Photo by R. Winter-bottom.

Fig. 12. Left lateral view of Trimma erdmanni, 10.2 mm SL juvenile paratype, ROM 85033, Kawe Island, Raja Ampat. Photoby R. Winterbottom.

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tial papilla and base of first anal-fin ray. Lateralscales 23-24 (x = 23.1, SD = 0.30, n = 20), ante-rior transverse scales 8-11 (x = 9.3, SD = 0.73, n =19), posterior transverse scales 8-9, (x = 8.1, SD =0.31, n = 19), no scales on cheek or opercle (except

18% length of fifth pelvic ray), first four rays withone or two sequential branch points, fifth raybranched once and 52-61-70% length of fourth (x= 64.5, SD = 4.44, n = 20), fourth ray reachingposteriorly to between anterior margin of urogen-

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trench. No dermal crest anterior to first dorsal fin.Epaxialis musculature extending anteriorly toabout a vertical with posterior margin of pupil(Fig. 7B). Abdominal/caudal vertebral pattern isType B, but haemal arch of second caudal vertebrais somewhat enlarged.

C o l o u r p a t t e r n (from slides of six freshlycollected specimens, 10.2-26.2 mm SL). A 26.2mm SL male (ROM 85033, Fig. 8) has an overallpinkish hue, darker on the dorsum, dominated bya slightly more than pupil-width midlateral orangestripe bordered with red, which continues anteri-orly onto the head behind the eye in a shallow “>”-shape, the lower limb touching the ventral marginof the eye, the upper limb to a point in line withthe dorsal margin of the pupil. In this specimen,the two arms of the “>” are joined across the inter-space by a narrow red-orange bridge, the interspaceis pinkish-grey. The red-orange colouration contin-ues onto the snout anterior to the orbit and theanterior part of both upper and lower lips. Thecheek is pale pinkish grey, bordered dorsally by athin (about one-fifth pupil-diameter in width)bluish-white stripe which ends at the vertical limbof the preopercle. The pectoral fin base and isth-mus are pale grey. A reddish-orange stripe curvesposterodorsally from the upper margin of the orbit,ending above the middle of the opercle. The napeis darker red than most of the rest of the body, andthe dorsal half of the caudal peduncle is somewhatless so, with the space beneath the dorsal fins palepink. Most of the scales on the dorsal half of thebody have a very thin margin of red chro-matophores along their posterior margins. Therows above the lateral scales and the row beneath ithave a wider band of darker red pigmentation. Theabdomen is pale pink, which grades into pale redposteriorly. The midlateral stripe is flanked by dif-fuse darker stripes about equal in width, formed byscattered dark chromatophores, which also under-lie the red stripe, especially in the abdominalregion. The elements of the first dorsal fin eachhave a yellow spot just distal to the base, and thereis a very diffuse, half-pupil diameter dark stripeabove the spots, with a scattering of melanophoresin the fin membrane above this. The anterior ele-ments of the second dorsal fin have similar yellowspots, and melanophores are densely scatteredthroughout the fin membrane. The central rays ofthe caudal fin have splotches of yellow a little dis-tal to their bases, with an overall rosy hue (espe-cially proximally and centrally), the membranes of

in two specimens with one or two cycloid scalesrespectively on upper part of opercle); midline ofpredorsal usually scaleless, but with 6 scales inholotype, 4 in a 20.9 mm SL female, and 6 scalesin front of first dorsal spine followed anteriorly bya naked area and then 2 more scales across midlinein a 26.2 mm SL male; usually 4 cycloid scales inposterior vertical row on pectoral base (once 5),with two vertical rows of scales anterior to this; 5-6-8 (x = 7.2, SD = 0.73, n = 20) prepelvic cycloidscales in midline anterior to pelvic-fin base; 12 cir-cumpeduncular scales; 8-9 (x = 8.1, SD = 0.31, n= 18) in midline between base of last anal ray andfirst procurrent caudal ray; body scales ctenoidexcept for cycloid scales on anterior belly midline,beneath and immediately posterior to pectoral-finbase, anterior scales on sides of nape, and alongbase of first dorsal fin; generally, body scales extendanteriorly on sides of nape to between a verticalabove anterior margin of opercle and just posteriorto posterior margin of eye (Fig. 7). Gill openingextending anteroventrally to a vertical below pupil.Upper jaw with outer row of curved, spaced,enlarged canines, decreasing in size posteriorly todistal tip of premaxilla, a band of several irregularrows of small conical teeth at symphysis, numberof rows decreasing posterolaterally and ending justposterior to bend of dentary, innermost rowslightly larger and directed posteriorly at symph-ysis, decreasing in size and becoming vertically ori-ented, and extending to distal tip of premaxilla.Lower jaw with an outer row of curved, spaced,enlarged canines ending at bend of dentary, severalrows of small, slightly curved conical teeth at sym-physis, grading to a single row at bend of dentaryand continuing to lower coronoid process as a sin-gle outer row, innermost row at bend of dentaryabruptly larger than other inner teeth (twice theheight) in males, decreasing somewhat in size pos-teriorly and ending at the upper margin of thecoronoid process. Cephalic sensory papillae as inTable I and Fig. 7. Tongue truncate with roundededges. Gill rakers on first arch 4-5 + 14-16-17 =19-21-22 (x = 20.4, SD = 0.73, n = 21), gill rakerswith obvious fleshy crenulations (Fig. 1A). Ante-rior nasal opening a short tube extending out overupper lip, posterior nasal opening a pore with araised rim, both protruding from slightly raisedoval sac confined to anterior half of snout. Bonyinterorbital 32-50-52% pupil width (x = 42.6, SD= 5.26, n = 20), with broadly U-shaped interorbitaltrench with sloping sides and no postorbital


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wedge over the hypural region and anterior bases ofthe caudal fin rays.

C o l o u r p a t t e r n i n a l c o h o l . The 26.2mm SL male is straw coloured, including all areasthat are red, orange or yellow in fresh specimens;however, most of the dark chromatophores and themelanophores remain obvious. The centre of thebody stripe above the pectoral fin has a diffuse pos-teriorly tapering darker stripe formed by scatteredsmall chromatophores, and the margins of themain stripe are bordered dorsally and ventrally bya dark diffuse pupil-width stripe of melanophoresthat tapers somewhat posteriorly and ends at thehypural plate. A few small melanophores are scat-tered over the body. The V-shaped anterior bifur-cation of the stripe is not apparent anteriorly, buteach arm of the bifurcation is bordered dorsally bya dark chromatophore band, and there is a narrowstripe of such chromatophores from the dorsalmargin of the pectoral fin base across the opercle tojust below the posterior margin of the eye. The redborders of the orange lateral stripe are more or lessdevoid of chromatophores, forming two palestripes bordered on either side by dark chro-matophores. The dorsal regions of the head andsnout have a scattering of melanophores, especiallydorsally. The interorbital trough has a longitudinaldark line about two chromatophores in width,which continues onto the snout as a variably widermedian stripe of melanophores from the anteriorpair of interorbital papillae almost to its anteriormargin. Other specimens are generally similar,with considerable variation in the degree of devel-opment of the dark chromatophores on the headand body. Comparisons: Trimma erdmanni is unique

among described species of the genus in itslive/freshly collected colour pattern of a red toorange body with a darker lateral stripe that usuallybifurcates anteriorly above the opercle, togetherwith a horizontal pale stripe from the upper jaw tothe opercle that abuts the ventral margin of theorbit. The species belongs to the grade possessing aType B configuration of the abdominal/caudal ver-tebral transition. Within this grade, subsequentsubdivision is usually made on the basis of presenceor absence of scales in the predorsal midline. How-ever, T. erdmanni, like a few large specimens of T.hayashii and a currently undescribed species fromCocos-Keeling, may have some scales in this region(although they are absent in most specimens, espe-cially when < 20 mm SL). The new species may be

the outer part of the fin appears greyish due tonumerous scattered melanophores. The anal finhas a faint pinkish hue proximally with manymelanophores scattered in the membranes distally.Pectoral and pelvic fin membranes are hyaline, therays of the median fins and the pectoral fin are palered. The iris is golden-yellow, with scattered redpigmentation, and heavily invested with dark pig-mentation (especially dorsally), with a thin whiteinner rim, and a dark blue band which crosses theiris from the anteroventral to the posterodorsalmargins. A live specimen (Fig. 9) is generally simi-lar, although the interspace of the “>” part of thelateral stripe on the head almost totally occluded,and the yellow spots in the dorsal fins are hardlydiscernable. The snout is orange, with the posteriornasal opening in a small white spot, a similar spotjust in fron of the eye, a median white stripe fromthe anterior interorbital region to the level of theposterior nares and another similar short stripefrom the posterior interorbital region over the dor-sum. In a 20.3 mm SL female (from the same lot),the mid-part of the lateral stripe is more yellowthan orange, and the yellow spots of the dorsal finsform stripes. A 23.5mm male (Fig. 10, holotype)has the whole body more heavily invested withmelanophores, especially anteriorly, and the pinksand oranges are replaced by yellow. The interspaceof the “>” on the side of the head is almostoccluded by the two arms of the stripe, the yellowspots in the first dorsal fin and anterior part of thesecond dorsal fin form diffuse stripes, the darkstripe above this is less discernable, but there is adiffuse dark stripe above the yellow in the seconddorsal fin; in both fins, there is a much greater con-centration of melanophores. The basal region ofthe anal fin is pale yellow rather than pink. A 14.3mm SL juvenile is very dark overall, with denselyscattered dark chromatophores on the head andbody (Fig. 11). The orange-red lateral stripe is onlydiscernable on the head and scarcely at all on thebody, and the yellow stripes in the dorsal fins aredistinct. A 10.2 mm SL juvenile (Fig. 12) has atranslucent body and head except for the darkchromatophores, which are accentuated along themidlateral orange-yellow stripe. The upper lip,snout, top of the head and “>” arms of the lateralstripe are yellow, and the median fins have a scat-tering of melanophores. A live specimen pho-tographed in Davao Gulf, Mindanao by ArthurBos has the posterior region of the lateral red stripeexpanded posteroventrally to a blunt triangular

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and blotches absent) – this species is apparentlyconfined to the Great Barrier Reef of Australia.Finally, T. tauroculum has an eye-diameter sizedblack ocellated blotch on the side of the bodyabove the pectoral fin, and numerous smaller blackspots on the nape (vs. both absent) – this specieshas only been recorded to date from Palau, andfrom Ulithi Atoll in the Yap Islands (Western Car-olines).Distribution: Trimma erdmanni is currently

known from the Raja Ampat islands and Sulawesiin Indonesia, and the Hermit Islands and Madang,Papua New Guinea, in water depths of 0-66 m.There are also photographs of a freshly collectedspecimen from El Nido, Palawan Island, and of livespecimens from Davao Gulf, Mindanao andCalamianes Is., Batangas in the Philippines (Fig.6). The species has also been observed in the Lem-beh Strait area of Sulawesi (G. R. Allen, pers.comm., shown with a “?” mark in Fig. 6).Etymology: The species is named for Mark V.

Erdmann in appreciation of his deep interest inTrimma (and other fishes, of course), his enthusi-astic collection and documentation of specimens ofthis genus for the present author’s research pro-gram, his friendship, and for the superb job hedoes for Conservation International’s IndonesianMarine Program. This species has been referred toinformally (in litt.) as Trimma RW sp. 68.

Trimma habrum n. sp.(Figs 6, 13-15)Delicate pygmy goby


Material Examined: A total of 1 lot, 12 type spec-imens, plus two additional non-type specimens(tissue voucher specimens). The description isbased on the holotype and 11 paratypes (15.5-17.7mm SL).Holotype: ROM 87486, 16.8 mm SL male,Indonesia, Raja Ampat, Fam Islands, Keruo Island(just east of Penemu Island), vertical wall,0°35’15.4”S; 130°17’41.1”E, 70 m, clove oil, 25January 2010, M. V. Erdmann.Paratypes: All specimens collected with the holo-type: AMS I.45592-00, 1, 17.5. MZB 19778, 4,15.5-17.6. ROM 84881, 5, 16.3-17.7. ROM1832CS, 1, 16.6 (male).Additional (Non-type) Material. Two specimens

distinguished from T. hayashii in having a shorterbasal membrane (< 20 vs. > 50% length of fifthpelvic fin ray) and usually more pectoral fin rays(17-19, x = 18.0 vs. 14-17 (x = 15.8) and from theundescribed species in having 8 vs. 9 anal fin rays.The predorsal scales, if present, are confined to anarea well anterior of the first dorsal fin spine andseparated from it by an unscaled area, in contradis-tinction to the other species possessing predorsalscales, in which the posteriormost scale lies imme-diately anterior to the spine. Further comparisonsare therefore confined to the species lacking scalesin the predorsal midline. Of the remaining 28described species without scales in the predorsalmidline, the presence of 19 or more total gill rak-ers in T. erdmanni separates it from all but eight ofthem (viz. T. benjamini Winterbottom, 1996, T.bisella Winterbottom, 2000, T. cana Winterbot-tom, 2004, T. capostriatum (Goren, 1981), T.necopinum (Whitley, 1959) T. sostra Winterbot-tom, 2004, T. striatum (Herre, 1945) and T. tauro -culum Winterbottom & Zur, 2007). Trimma capo -striatum and T. striatum can be immediately sepa-rated by the presence of six thin red stripes on thehead and anterior part of the body, and in possess-ing a full basal membrane and 2-3 branches in thefifth pelvic fin ray (vs. no thin red stripes confinedto the head and anterior body, basal membrane 7-18% the length of the fifth ray, which is branchedonly once). Trimma benjamini is most easily sepa-rated from the new species by the thin ring ofmelanophores encircling the eye, with two short,ventrally directed bars onto the cheek (vs. suchmarkings absent) and in lacking the dark lateralstripe on the body that usually bifurcates behindthe eye. The two species are very similar to eachother in their meristic values. Trimma bisella usu-ally has 10 dorsal and 9 anal fin rays (vs. 9 and 8respectively), and has an unbranched fifth pelvicfin ray, has a large white blotch on the dorsal sur-face of the caudal peduncle (vs. blotch absent) andfour orange bars on the head (vs. bars absent) – thisspecies is apparently confined to the westernIndian Ocean (Mauritius). Trimma cana and T. sos-tra have an unbranched fifth pelvic fin ray (vs.branched) and a translucent to white body withnumerous red bars (T. cana) or blotches (T. sostra)in life (vs. such red markings absent). Trimmanecopinum usually has 9 anal rays (vs. 8), a fullbasal membrane (vs. much reduced), and alive/fresh colour pattern of numerous red to orangespots and blotches on the head and body (vs. spots

Fig. 13. Dorsal view of the head of Trimma habrum (16.8mm SL male holotype, ROM 87486). Specimen stainedwith cyanine blue. Photo by R. Winterbottom.

Fig. 14. Left lateral view of the vertebral transition ofTrimma habrum, 19.2 male paratype, ROM 1832CS.Specimen stained with alizarin and alcian blue. Photo byR. Winterbottom.

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tical line above a line between urogenital papillaand anal spine; pelvic fin I 5, no frenum, basalmembrane absent, first four rays with one sequen-tial branch, fifth ray unbranched and 44-46-52%length of fourth (x = 46.8, SD = 2.88, n = 11),which reaches posteriorly to between bases of firstto third anal-fin ray. Lateral scales 23 (n = 11),anterior transverse scales 7-8 (x = 7.2, SD = 0.25),posterior transverse scales 7-7.5 (x = 7.0, SD =0.14), scales across predorsal midline 9-10 (x = 9.2,SD = 0.37), cheek with 1-2 scales in upper row and7-8 cycloid scales in main lower row, upper two-thirds of opercle covered with three longitudinalrows of 2, 2 and 1 cycloid scales respectively (theposterodorsal scale may occasionally be weaklyctenoid); 2 vertical rows of cycloid scales on pec-toral base, with 3 scales in posterior row; 4 cycloidprepelvic scales in midline; 11-12 (x =11.7, SD =0.47) circumpeduncular scales; body scales ctenoidexcept for cycloid scales on anterior belly midline,beneath and immediately posterior to pectoral-finbase, and first few scales in predorsal midline andaround posterodorsal margin of eye (Fig. 13). Gillopening extending anteroventrally to a verticalbelow anterior third to middle of pupil. Upper jawwith an outer row of somewhat enlarged, spaced,slightly curved canines decreasing in size posteri-orly to end of premaxilla, 2-3 irregular inner rowsof small conical teeth behind symphysis decreasingto a single inner row extending to end of premax-illa; innermost row slightly larger and directed pos-teriorly, decreasing in size posteriorly. Lower jawwith an outer row of enlarged curved spacedcanines from symphysis to bend of dentary anddirected somewhat anterodorsally, 1-2 irregular

collected with ROM 84881 for genetic analysis:ROM T07710 and T07711 (17.3-18.2).Diagnosis: Trimma habrum is characterized by a

bony interorbital as wide as the diameter of thepupil, 8-9 scales in the predorsal midline, 14unbranched pectoral fin rays, an unbranched fifthpelvic fin ray, no basal membrane joining the fifthpelvic fin rays across the midline, a single full rowof cheek scales and sometimes one or two smallscales above this row, three rows of scales coveringthe upper two-thirds of the opercle, and no trace ofdark or black pigmentation on the caudal pedunclewhen preserved. The fresh colouration is also diag-nostic: a pale translucent dorsum with light yellowblotches and the base of each element of the dorsalfin surrounded by a red spot, a thin red bar alongthe posterior margins of the hypurals, no dark pig-ment at all on the hypural region of the peduncle,and a darkly pigmented covering to the dorsal mar-gins of the abdominal cavity and the brain.Description: Dorsal fins VI + I 8-9 (x = 8.8, SD

= 0.37), second and third spines longest, secondspine reaching to between base of second ray ofsecond dorsal fin and one scale posterior to last raywhen adpressed (to base of sixth ray on holotype),third to between base of spine and first ray of sec-ond dorsal fin, last ray extends posteriorly forabout one-third of distance between its base andfirst dorsal procurrent ray, all rays branched except,usually, posterior element of last ray; anal fin I 8-9(x = 8.1, SD = 0.28), all rays branched except pos-terior element of last ray, which extends posteriorlyfor about one-third of distance between its baseand first ventral procurrent ray; pectoral fin 14, allrays unbranched, fin reaching posteriorly to a ver-

Fig. 15. Left lateral view of Trimma habrum, 16.8 mm SL male holotype, ROM 87486, Keruo Island, Raja Ampat. Photoby R. Winterbottom.

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the base of the anal fin and ending about at theanterior third of the peduncle. The whole ventralhalf of the body behind the abdominal cavity isreddish-pink. The snout is yellow with a reddishnasal sac, the brain case is heavily sprinkled withlarge dark pigment cells, and the cheek has a whitestripe between the posteroventral and anteroventralmargins of the eye. Below this, the cheek is yellowgrading to reddish-pink posterodorsally; the oper-cle has a light mauve suffusion. The pectoral finbase is suffused with light purplish-red, the base ofthe pelvic fin is suffused with yellow. The spines ofthe first dorsal fin are red, each with a slightlyenlarged red spot at its base, and with a yelloworange, half-pupil diameter wide yellowish orangestripe just above the base. The membranes betweenthe first three spines are heavily sprinkled withmelanophores, the fin membrane posterior to thisis hyaline. The bases of all elements of the seconddorsal fin are contained in red spots, the proximalhalf of the fin membranes is heavily suffused withyellow (more so posteriorly), and the distal halfcontains scattered melanophores and red and yel-low chromatophores. The caudal fin contains a mixof yellow and red suffusions with scatteredmelanophores, and the bases of the rays and theregions adjacent to them are red, forming a thinred vertical line over the ends of the hypurals. Thebasal region of the anal fin has a series of redinverted triangle separated by pale areas (these donot conform to the bases of the fin elements), thefin has a basal black stripe followed by a broad yel-low stripe and culminating in a distal black stripe.The pectoral fin rays are red, intensified at theirbases, and the membrane is hyaline. The pelvic fin

inner rows of smaller curved teeth at symphysisgrading to a single row laterally and continuing tomid-height of coronoid process of dentary.Cephalic sensory papilla counts as in Table I.Tongue broadly truncate with rounded edges. Gillrakers on first arch 3-4 + 12-13 = 15-16-17 (x =3.1 + 12.9 = 16.0, SD = 0.28, 0.28, and 0.41respectively). Anterior nasal opening a short tubeextending out over upper lip, posterior nasal open-ing a pore with a raised rim, both protruding fromslightly raised oval sac confined to anterior half ofsnout. Bony interorbital equal to pupil width, witha gently rounded fleshy median interorbital ridge,no dermal crest, and epaxialis musculature extend-ing anteriorly to above a vertical with posteriormargin of pupil (Fig. 13). Abdominal/caudal verte-bral configuration Type A, with haemal arches offirst two caudal vertebrae expanded to accommo-date posterior extension of swimbladder (Fig. 14).

C o l o u r p a t t e r n (from slides of two speci-mens, 16.8-17.3 mm SL, description based mostlyon the former, Fig. 15). The 16.8 mm SL maleholotype is a essentially a pale, translucent fish,with the anterior region of the dorsum suffusedwith light yellow, the exposed portions of the scalesbelow the dorsal fins are thinly edged with red,there are two yellow saddles a little less than pupil-diameter in width over the dorsal part of thepeduncle, and one (below the second dorsal saddle)over the ventral part of the peduncle. The neuralcanal and upper half of the swimbladder are heav-ily invested with melanophores and clearly visiblein lateral view; the abdomen is pale pink anteriorlygrading to yellow posteriorly, the yellow intensify-ing into a narrow, tapering wedge posteriorly above

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Trimma haimassum n. sp.(Figs 1 B & C, 2A, 6, 16-20)Blood-spot pygmy goby


Trimma sp. – Hayashi & Shiratori 2003: 44 (#070), 45 (# 078); Indonesia)

Trimma sp. 1 – Kimura & Matsuura 2003: 193(Bitung, Indonesia)

Trimma sp. 1 (R. Winterbottom sp. 63) – Allen &Erdmann 2009: 619 (Fakfak/Kaimana and RajaAmpat, Indonesia)

Trimma sp. 10 – Kuiter & Tonozuka 2004: 706(Indonesia)

Material Examined: A total of 11 collections, 240(10.4-30.9 mm SL) type specimens, plus addi-tional non-type specimens from Raja Ampat(including a tissue voucher specimen), as well asnumerous specimens from several other westernPacific localities. The morphological description isbased primarily on 10 males, 9 females (25.0-30.1mm SL, x = 27.6, SD = 1.52, ROM 85347), plusthe holotype, with some details (teeth, vertebraltransition) from ROM 1836CS (5, 19.5-27.3); allspecimens from the same collection.Holotype: ROM 87484, 27.3 mm SL female, Eside of “Barracuda Rock”, 200 m N of Wayil Island(02°11’43.4”S, 130°25’37.9”E), cave with lots ofsea fans & sea whips near entrance at top part, bigvariety of soft & hard corals just outside, 19.8-27.4m, rotenone, 1300-1340, field # RW10-40, 1 Feb-ruary 2010, R. Winterbottom, L. Katz & CI team. Paratypes: AMS I.45593-001 (formerly ROM85390), 38, 14.6-30.9, SE islands off Misool,south side of Balbulol Island (02°01’29.5”S,130°41’34.9”E), 45 m, rotenone, 0750-0810, field# RW10-45, 2 February 2010, M. V. Erdmann.MZB 19779 (formerly ROM 85377), 59, 14.1-27.7, SE islands off Misool, south side of BalbulolIsland (02°01’29.5”S, 130°41’34.9”E), sea fans,sea whips, sponges, tunicates, hydroids, 13.7-18.3m, rotenone, 0750-0850, field # RW10-44, 2 Feb-ruary 2010, R. Winterbottom, L. Katz & CI team.ROM 84877, 5, 21.0-27.5, Keruo Island, off Pen-emu Island, Fam Islands (00°35’15.4”S,130°17’41.1”E), various hard and soft corals, tuni-cates and sponges, 18.3-27.4 m, clove oil, 0810-0850, field # RW10-03, 25 January 2010, R. Win-terbottom & L. Katz. ROM 84884, 2, 20.4-21.8,Keruo Island off Penemu Island, Fam Islands

and fin rays are hyaline. The iris is yellow with adiffuse black margin and a horizontal, one-thirdpupil-diameter dark purple stripe which touchesthe dorsal rim of the pupil ventrally. The secondphotographed specimen (17.3 mm SL, ROMT7710, tissue voucher) is essentially similar, butlacks the yellow wedge from the abdomen thatpasses posteriorly above the base of the anal fin.

C o l o u r p a t t e r n i n a l c o h o l : off white,with darker pigmentation over the swimbladdervisible through the body, as well as that coveringthe braincase around the posterior rim of the orbit.The dark pigmentation in the distal portions of thedorsal fins, the base and distal regions of the analfin, and a few dark chromatophores on the dorsalsurface of the snout are also visible.Comparisons: Trimma habrum belongs to the T.

tevegae species complex, which is defined by theapparently apomorphic conditions of the broadbony interorbital (75% or more of pupil diameter)and a Type A abdominal/caudal vertebral transi-tion region. Among the 10 nominal species cur-rently assigned to this group, T. habrum differsfrom T. caudomaculatum Yoshino & Araga, 1975,T. griffithsi Winterbottom, 1984, T. nasa Winter-bottom, 2005 and T. tevegae Cohen & Davis,1969, as well as from T. xanthochrum n. sp.described herein, in lacking any dark pigmentationon the posterior portion of the caudal peduncle.All the remaining species except T. marinae Win-terbottom, 2005, have at least some branched raysin the middle of the pectoral fin. Trimma marinaeshares with T. habrum the thin vertical red bar overthe ends of the hypurals in freshly collected speci-mens, but has an open nasal capsule lacking ante-rior and posterior nares (vs. a nasal sac with bothnares present), fewer scales in the predorsal midline(6-8 vs. 9-10), and the bases of the elements of thedorsal fin are not surrounded by a small, dark redspot (vs. such a spot present).Distribution: Trimma habrum is currently only

known from a single collection at 70 m on a verti-cal wall at Keruo Island (off Penemu Island), one ofthe Fam Islands in Raja Ampat, Indonesia (Fig. 6).Etymology: Derived from the Greek word

‘habros’, meaning delicate, dainty or graceful, inallusion to the soft and delicate shades of colour offreshly collected specimens. Trimma habrum hasbeen referred to informally (in litt.) as T. RW sp.95.

Fig. 16. Left lateral view of accessory scales above end ofpectoral fin of Trimma haimassum (27.3 female paratype,ROM 85347). Specimen stained with cyanine blue. Photoby R. Winterbottom.

Fig. 17A-B. Trimma haimassum, 27.3 female paratype,ROM 85347, Wayil Island, Raja Ampat. Left lateral (A)and dorsal (B) views of head to show papillae, specimenstained with cyanine blue. Photo by R. Winterbottom.


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(00°35’15.4”S, 130°17’41.1”E), 70 m, clove oil,0815-0850, field # RW10-04, 25 January 2010,M. V. Erdmann. ROM 84895, 6, 15.4-24.5,Keruo Island off Penemu Island, Fam Islands(00°35’15.6”S, 130°17’41.1”E), 56 m, clove oil,1200-1300, field # RW10-05, 25 January 2010,M. V. Erdmann. ROM 85035, 1, 22.0, KaweIsland, SW bay (00°05’15.2”S, 130°07’25.3”E),50 m, clove oil, field # RW10-10, 26 January2010, M. V. Erdmann. ROM 85084, 1, 15.5,about 1.2 km SSE of Mutus Island, on west side ofsand spit (00°21’01.0”S, 130°21’25.4:E), sponges,tunicates, some hard and soft coral, 50 m, clove oil,0900-0920 field # RW10-16, 28 January 2010, M.V. Erdmann. ROM 85156, 20, 14.3-25.8, TanjungManare, Waigeo Island, west side at about middleof width off a small cape (00°16’26.1”S,130°19’01.5”E), hard and soft corals, sponges,19.8-22.9 m, rotenone, 0750-0820, field # RW10-23, 29 January 2010, R. Winterbottom, L. Katz, P.Johannes, W. Kaka & W. Awom. ROM 85347, 66,16.6-30.0, collected with the holotype. ROM87415, 1, 16.4, Ef Pian Island (SE of Misool),south side (02°02’28.8”S, 130°46’41.5”E), 45 m,clove oil, 1530-1550, field # RW10-48, 2 February2010, M. V. Erdmann. ROM 1836CS, 5, 19.5-27.3, collected with the holotype. USNM 399287(formerly ROM 85210), 35, 10.4-24.9, WofohIsland, west coast near south end (00°15’21.9”S,130°17’32.0”E), many small corals, Tubastrea, seawhips, some black coral, sea fans, tunicates,sponges, some hydroids, 18.3-22.9 m, clove oil,1500-1540, 29 January 2010, R. Winterbottom &W. Kaka.Additional (Non-type) Material. Indonesia: Flo-

res, BPBM 36697, 2 (21-26). Raja Ampat, ROMT07712 (24), collected with ROM 84884, forgenetic analysis. ROM 84905, 6 (12.7-21.3);ROM 85219, ( 25.2); ROM 85226, 4 (20.7-23.9);ROM 85317, 3 (10.2-10.8); ROM 85328, 2(16.0-21.8); ROM 87408, 5 (13.0-29.7); ROM85106, 3 (11.7-21.4); ROM 85137, 6 (16.0-20.6);ROM 85147, 4 (11.1-23.5); ROM 87424, 7(18.2-28.4); ROM 87434, 8 (16.0-26.9); ROM87452, 5 (14.4-22.1); ROM 87477, 6 (16.0-25.4).Saparua, USNM 210092, (17); USNM 209987,(20). Sulawesi, Ujung Padang, ex-BPBM 26808, 3(20-23). Papua New Guinea: Bougainville Island,WAM P.28161-011, 5 (12-24). Hermit Is., JalunIsland, USNM 264557, (22); Madehas Island,WAM P.28165-011, (18). Manus Island, ex-WAMP.27826-086, (18). New Britain, Rabaul, ex-WAM

P.28172-041, (25); WAM P.28174-025, 5 (17-29).Philippines: Cebu, ROM 1137CS, 3 (19-30);ROM 54753, 8 (19-30). Negros Oriental, ROM54742, 6 (22-26). Palawan, El Nido, BLIH1983354, 3 (25-28). Siquijor Island, ROM 54741,(23); ROM 69534, (24); ROM 69583, (17).Solomon Islands: Guadalcanal, ROM 54754,(23); ROM 54755, 2 (15-22). Diagnosis: Trimma haimassum is characterized by

a relatively deep body, numerous irregular scaleswith about 30 lateral rows and over 15 scales in theanterior transverse series, a fifth pelvic fin ray thatbranches twice dichotomously, usually a somewhatelongate second dorsal spine reaching to a littlebeyond the spine of the second dorsal fin, no scaleson the midline of the predorsal region, cheek oropercle, a moderately wide bony interorbital that is

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Fig. 18. Left lateral view of Trimma haimassum, 25.9 female paratype, ROM 85347, Wayil Island, Raja Ampat. Photo by R.Winterbottom.

Fig. 19. Left lateral view of Trimma haimassum, 27.5 mm SL male paratype, ROM 84877, Penemu Island, Raja Ampat.Photo by R. Winterbottom.

about 55% the width of the pupil and with a fleshymedian ridge between the eyes, and a dermal ridgeanterior to the first dorsal spine. The red spotabove the opercle, which continues a thin red linebelow the orbit where it is underlain by a bluestripe, is distinctive in live individuals. In fresh andpreserved material, the red spot and line are dark,there are two dark stripes between the anteroven-tral margin of the eye and the edge of the upper lip,and the tip of the snout has a dark reticulated pat-tern, with dark transverse stripes over the dorsalmargin of the orbit.Description: The number of specimens (n) is

given only where they are less than 20. The bodyscales in the species are in numerous, irregular rowsthat may be partially incomplete, and accessoryscales are usually present (Fig. 16), making scale

counts exceedingly difficult and potentially unreli-able. Dorsal fins VI + I 10, second spine longest,usually reaching to between bases of first to fourthrays of second dorsal fin when adpressed (not elon-gate in two males, 29.6-29.9 mm SL, reachingeither to base of spine of second dorsal fin or inter-space between fins), all rays branched except, usu-ally, posterior element of last ray; anal fin I 8-9-10(x = 9.0, SD = 0.37), all but first and posterior ele-ment of posteriormost ray usually branched (firstelement branched in four specimens); posterior-most ray of both fins reaching between half andthree-quarters of distance between ray base andfirst dorsal or ventral procurrent ray; pectoral fin18-19 (x = 18.5, SD = 0.50), 3-4 (x = 3.2, SD =0.39) and 1-3-4 (x = 2.5, SD = 0.66) unbrancheddorsal and ventral rays respectively, middle raysbranched, fin reaching posteriorly to a vertical linebetween middle of urogenital papilla and base ofanal spine; pelvic fin I 5, no frenum, basal mem-brane 25-55% length of fourth ray (n = 15, highvariance probably due to delicate nature of mem-brane, which is easily torn), first four rays with 2-4sequential branch points, fifth ray branchedbranched twice dichotomously and 59-68-76%length of fourth (x = 67.6, SD = 4.51), fourth rayreaching posteriorly to between urogenital papillaand base of second anal fin ray. Although seg-mented caudal fin rays nearly always total 17 (1 of18 specimens with 16), variation in number ofbranched vs. unbranched rays was found; dorsalhalf usually with 3 unbranched and 6 branchedrays (once 2/6, once 1/7); ventral half usually with5 branched, 3 unbranched (n = 10) or 6 branched,

2 unbranched (n = 7), once 7/2. Lateral scalesabout 29-31 (x = 30.3, SD = 0.63), anterior trans-verse scales about 15-16-17 (x = 16.7, SD = 0.83),posterior transverse scales about 10-12-13(x =11.6, SD = 0.79), no scales in midline of predorsal,cheek or on opercle; pectoral base with 4-5 verticalrows of cycloid scales, outer row of 7-8-10 (n = 16)scales, anteriormost row largest, posteriormostsmallest; 7-9 prepelvic cycloid scales in midlineanterior to pelvic-fin base (2 specimens with 11,where scales extended anterior to ventral tips ofcleithra on isthmus); 16-18 circumpeduncularscales (x = 16.9, SD = 0.70, n = 18); body scalesctenoid, extending from above posterior third ofopercle, passing slightly anteriorly and then up andback in an arc, with a narrow scaleless border alongbase of first dorsal fin up to about base of 5th ray;scales cycloid anterior to a line about 3 scales pos-terior to axil of middle of pectoral base to aboutmid region of belly. Gill opening extendinganteroventrally to a vertical below anterior third tomiddle of pupil. Upper jaw with outer row ofspaced, curved, enlarged canines, decreasing some-what in size posteriorly and extending to end ofpremaxilla, 3-4 irregular rows of small conical teethbehind the outer row at symphysis, those near sym-physis in innermost row directed posteriorly,enlarged, but decreasing to size of other inner rowteeth posterolaterally, which grade to single rowposteriorly at end of premaxilla. Lower jaw withouter row of spaced, curved, enlarged canines, end-ing at bend of dentary where these teeth are largest,several irregular rows of small conical teeth at sym-physis grading to single outer row ending at

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Fig. 20. Left lateral view of Trimma haimassum (live), Misool Island, Raja Ampat, Photo by G. R. Allen.

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distal to the base, and the fin membranes are heav-ily sprinkled with melanophores. The caudal finhas scattered, half-pupil diameter sized yellow toorange spots and numerous melanophores in thefin membranes. The anal fin also has numerousmelanophores in the membranes, and is suffusedwith reddish-pink. The pelvic fin is pinkish, thepectoral fin has reddish fin rays and hyaline mem-branes. A 27.5 mm SL male (Fig. 19) has a moreorange head and body, the blue lines under and infront of the eye are intermittent or absent as is thered stripe between the ovoid spot above the end ofthe opercle and the opercle, the basal stripe in thedorsal fins is vivid orange, there is a line of orangespots in the second dorsal at about the middle oftheir length, the caudal spots are orange, as is thebasal half of the anal fin, and the scale pockets onthe dorsal half of the body are more heavily out-lined with dark chromatophores. A specimen pho-tographed underwater (Fig. 20) has a much yel-lower head, the red stripe from the ovoid opercularspot is in two parts, and the bars across the orbitand lines on the snout are blue. An anaethetisedphotographed underwater has the red opercularblotch continuing anteriorly across the ventralmargin of the pupil almost to the maxilla, and theblue line beneath it extends posteriorly to the mid-region of the opercle. The area immediately abovethis red stripe on the snout is bright yellow, and thethin blue stripe above it is margined with red. Thediagonal stripe across the iris is mauve, as are theshort stripes to the midline on the fleshy part of theanterodorsal orbit. The median fins are moregreenish-yellow, but this may be due to the darkblue background, and the scale pockets on the dor-sum are relatively strongly outlined. The remainingphotographed specimens are similar overall,although some have many more dark pigment cellsin the fin membranes (including the pelvic fin),may have a more or less distinct dark basal band inthe dorsal fins, and the overall body colour may bebrown rather than red, pink or yellow.

C o l o u r p a t t e r n i n a l c o h o l : head andbody pale straw-yellow; the nape, upper half of theopercle and the body, except for the anteroventralregion of the abdomen, with a dense scattering ofmelanophores and subdermal brown chro-matophores, which tend to be concentrated aroundthe scale pockets anterodorsally in specimens inwhich the scales have been abraded off. An ill-defined, irregular line primarily of mela no phoresruns anteroventrally from the anterodorsal region of

anterodorsal tip of coronoid process of dentary,innermost row somewhat enlarged (half the size ofouter teeth), spaced, and continuing posteriorly tobeginning of coronoid process. Cephalic sensorypapillae as in Fig. 17 and Table I (nomenclature ofpapillae given in Fig. 2A). Tongue truncate withrounded edges. Gill rakers on first arch 4-5 (x =4.4, SD = 0.49) + 16-17-18 (x = 16.9, SD = 0.71)= 20-21-22 (x = 21.3, SD = 0.71), larger rakers onboth epibranchial and ceratobranchial possess arow of small, bluntly pointed crenulations alongtheir medial surfaces (Fig. 1B). Anterior nasalopening a short tube extending out over upper lip,posterior nasal opening a pore with a raised rim,both protruding from slightly raised oval sac con-fined to anterior half of snout, posterior openingabout 2-3 X its diameter from closest border of eye.Bony interorbital 47-64% pupil width (x = 55.5,SD = 5.07), interorbital profile broadly U-shapedwith a central ridge of soft tissue separating the leftand right sensory papillae row p (Fig. 17 B). Ridgeof tissue (dermal crest) extending from proximalone-tenth of first dorsal spine anteriorly, becomingreduced in height anteriorly and fading out aboveanterior third opercle. Epaxialis musculatureextending anteriorly to above a vertical with poste-rior third of pupil. Abdominal/caudal vertebraetransition Type B.

C o l o u r p a t t e r n (based on images of ananaesthetised specimen and nine freshly collectedspecimens). A 25.9 mm SL gravid female (Fig. 18)has a reddish head, orange-red dorsum grading toyellowish pink posteriorly, slightly lighter over theabdomen, with some of the more dorsal scale mar-gins slightly darker. An ovoid red spot, aboutpupil-diameter in width lies on and above the pos-terodorsal margin of the opercle and is connectedto the posteroventral margin of the orbit by anintermittent red stripe. A thin blue line underliesthe anterior portion of this stripe and continuesanteroventrally, skirting the ventral margin of theorbit and ends just above the upper jaw. A shortblue line from the orbit to just above the middlelength of the maxilla is present just dorsal to thisline, and is similarly oriented. Four short blue linesbetween the anterior and dorsal margins of thefleshy orbit pass medially towards the midline. Theiris is red and speckled with darker pigment, andthere is a thin blue line passing across it fromanteroventral to posterodorsal oriented at about45°. The dorsal fins have an orange stripe, aboutone-third pupil diameter in width, situated just


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opercular margin anteriorly to the posterior marginof the eye (vs. absent). Distribution: Trimma haimassum has been col-

lected from south-western Sulawesi north toPalawan in the Philippines and eastwards to theSolomon Islands (Fig. 6) in depths of 7-70 m. Itmay also occur in Palau. This was the most abun-dant species of the genus at Raja Ampat.Etymology: Derived from the Greek word

‘haimasso, stained with blood, in allusion to theblood-red spot above the posterior end of the oper-cle in live and fresh specimens. This species hasbeen referred to informally (in litt.) as Trimma RWsp. 63.

Trimma papayum n. sp.(Figs. 6, 21-24)Pawpaw pygmy goby


Trimma sp 6 – Kuiter & Tonozuki 2004: 705 (twounderwater photographs, Maumere, Flores)

Material Examined: A total of 2 lots, 12 type spec-imens, plus one additional non-type specimen (tis-sue voucher). The description is based on the holo-type and up to 11 paratypes (11.5-22.2 mm SL).Holotype: ROM 87485, 21.6 mm SL female,Indonesia, Raja Ampat, SW bay of Kawe Island,00°05’15.2”S, 130°07’25.3”E, 50 m, clove oil, 26January 2010, M. V. Erdmann.Paratypes: AMS I.45594-001, (20.5), collectedwith the holotype. MZB 19780, 4 (16.8-20.5),collected with ROM 87564. ROM 85034, 2,12.2-21.1 mm SL, collected with the holotype.ROM 1833CS, 19.7 mm SL, collected with theholotype. ROM 87564, 3 (11.5-22.2), Indonesia,Raja Ampat, SW Kawe Island, 00°05’46.6”S, 130°07’07.2”E, 50 m, clove oil. M. V. Erdmann, 28August 2010.Additional (Non-type) Material. One specimen

for genetic analysis, ROM T07725, (20.5), col-lected with the holotype.Diagnosis: Trimma papayum is unique among the

described species of the genus in having a one-thirdpupil diameter black spot on and just behind thefourth dorsal fin spine. It has 9 dorsal and 8 analfin rays, 18-19 pectoral rays with the middle onesbranched, a single branch point in the fifth pelvicfin ray, which is about 60-70% the length of thefourth, and a basal membrane about 10-20% the

the opercle across the ventral margin of the orbit andends above the maxilla; a similar short stripe liesabove this and passes parallel to it from theanteroventral margin of the orbit to above the mid-dle of the maxilla. The snout and both lips have anirregular reticulated mottling of melano phores. Thedorsal margin of the orbit has four to five transversestripes of melanophores. The cheek and pectoral finbase have a few scattered melanophores, and thedorsal margin of the pectoral fin base occasionallyhas a thin dark stripe; the membranes of theunpaired fins are heavily invested with melano -phores, while those of the paired fins have only a fewwidely scattered mela no phores. There is little evi-dence of the sub-basal stripe in the dorsal fins pre-sent in freshly collected specimens, although somespecimens do seem to have fewer melano phores inthe region where the stripe occurs.Comparisons: The new species belongs to the T.

sheppardi species complex, a group which lacksmedian predorsal, cheek and opercular scales, pos-sesses a deep head and body (body depth at base ofpelvic spine about 30% SL), has accessory scalesand disjunct scale rows, has a well developed nasalapparatus occupying the anterior half of the snoutwith the posterior opening in a short tube, hasmedian fleshy ridges in the interorbital region andpossesses a dermal crest on the nape just anterior tothe origin of the first dorsal fin. Trimma haimassumdiffers from all the other species in the group bynearly always having an elongate second dorsal finspine (vs. rarely elongate), having two (vs. one)dichotomous branch points in the fifth pelvic finray, in apparently in having more lateral scale rows(29-31 vs. 24-26 – uncertainty due to the difficultyin accurately recording this count), in the darkreticulations on the dorsal surfaces of the snout andlips, in having two dark stripes between the eye andthe maxilla, and in the colouration of the dark spotabove the opercle in life (red vs. black). It differsfurther from T. sheppardi Winterbottom, 1984, inhaving 10 (vs. 8-9) dorsal-fin rays; more outer gillrakers on the first gill arch (usually 4-5 + 15-18 vs.2-3 + 13-15); in lacking the two yellow bars on thecheek beneath the eye; and in having the anteriorportion of the head orange-red rather than pink. Itdiffers from T. yanoi Suzuki and Senou, 2008, inlacking the obvious internal dark blotches alongthe vertebral column that are especially prominentin live and freshly collected material. It differs fur-ther from an undescribed species (T. RW sp. 24) inhaving the extension of the dark areas above the

of 3 scales on the upper margin of the opercle, anda shallow interorbital trench with no trench pos-terodorsal to the orbit. Freshly collected specimensare orange red in overall colouration, with scattereddiffuse yellow spots.Description: Dorsal fins VI + I 9, second and

third spines longest, second reaching to betweenbase of fifth ray of second dorsal fin and two scalesbeyond end of that fin when adpressed, thirdreaching to bases of first to fourth ray of seconddorsal fin, rays all branched except posterior ele-ment of last ray, last ray reaching to between 55-60-90% distance from its base to first dorsalprocurrent fin ray (x = 67.1, SD = 10.10, n = 12) ;anal fin I 8, all but first and posterior element ofposteriormost ray branched; posteriormost rayreaching between 50-75% (x = 58.7, SD = 7.59, n= 12) of distance from its base to first ventralprocurrent fin ray; pectoral fin 18-19 (x = 18.4, SD= 0.49, n = 12), 3-4 dorsal and 3-6 ventralunbranched rays (but 6 and 9 respectively in the11.5 mm SL juvenile), middle rays branched, finreaching posteriorly to between a vertical lineabove urogenital papilla and base of first ray of analfin; pelvic fin I 5, no frenum, basal membrane 9-18% length of fifth ray, first four rays with onesequential branch (once with two branch points infourth ray), fifth ray branched once dichotomouslyand 56-65-74% (x = 64.8, SD = 5.15, n = 10)length of fourth, fourth ray reaching posteriorly tobetween bases of first to fifth anal-fin rays. Lateralscales 23 (n = 11); anterior transverse scales 9-11 (x= 10.1, SD = 0.69, n = 11); posterior transversescales 8-9 (x = 8.5, SD = 0.50, n =11); 5-6-10 (x =7.1, SD = 1.66, n = 9) scales in midline of predor-sal (fewest in smaller specimens, absent in 11.5mm SL specimen); no scales on cheek; a single rowof 3 cycloid or weakly ctenoid scales on opercle; 3-4 vertical rows of scales on pectoral fin base with4-5 cycloid scales in posterior row; 5-8 (x = 6.5, SD= 0.78, n = 11) cycloid prepelvic scales; 12 circum-peduncular scales; 8 rows of scales in midlinebetween base of last anal ray and first procurrentcaudal ray; body scales ctenoid except for cycloidscales on anterior belly midline, breast, beneathand just posterior to pectoral-fin; body scalesextend anteriorly to vertical in line with posteriormargin of orbit. Gill opening extending anteroven-trally to a vertical below middle to anterior one-third of pupil. Upper jaw with outer row of slightlycurved, enlarged canines decreasing in size posteri-orly to end of premaxilla, 2-3 irregular inner rows

Fig. 21A-C. Left lateral (A), dorsal (B), and ventral (C)views of head to show head papillae of Trimma papayum(19.6 mm SL male paratype, ROM 87564). Specimenstained with cyanine blue. Photo by R. Winterbottom.

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length of the fourth ray, 10-11 anterior and 8-9posterior transverse scale rows, 5-10 scales in thepredorsal midline, no cheek scales and a single row

of small conical teeth near symphysis decreasing toa single row to end of premaxilla. Lower jaw as forupper jaw, but outer row ends at bend of dentary,inner row enlarged at symphysis and decreases insize posteriorly to end at coronoid process.Cephalic sensory papillae as in Fig. 21 and Table I.Tongue truncate with rounded margins. Gill rakerson first arch 4-5 (x = 4.4, SD = 0.48) + 14-16 (x =14.6, SD = 0.77) = 18-20 (x = 19.0, SD = 0.95, n= 11). Anterior nasal opening a short broad tubeextending out over upper lip, posterior nasal open-ing a pore with a raised rim, both protruding fromslightly raised oval sac, posterior pore about 1 pore-diameter from nearest point of bony orbital rim.Bony interorbital 22-40-47 (x = 33.7, SD = 6.91)% pupil width, with broad shallow U-shapedinterorbital trough and no postorbital trough ortrench. No or very slight dermal crest anterior tofirst dorsal fin, epaxialis musculature reaching

Fig. 22. Left lateral view of Trimma papayum, 21.6 mm SL female holotype, ROM 87485, Kawe Island, Raja Ampat. Photoby R. Winterbottom.

Fig. 23. Left lateral view of the first dorsal fin to show thesize and position of the dark ocellated spot of Trimmapapayum (21.1 mm SL female paratype - preserved, ROM85034). Note damaged second dorsal spine. Photo by R.Winterbottom.

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almost to a vertical above posterior margin of orbit.Abdominal/caudal vertebral transition Type B.

C o l o u r p a t t e r n (from a freshly collectedspecimen, 21.6mm SL female holotype, Fig. 22).Overall orange-red (red especially evident alongthe middle of the trunk, above the anal fin and onthe peduncle), with numerous diffuse pupil-diam-eter yellow spots on the body at the centres of thescales, decreasing somewhat in size both anteriorlyon the nape and posteriorly on the peduncle, thoseon the nape yellow-orange rimmed with red, therest of the nape and the area immediately below thedorsal fins grey with the exposed margins of thescales diffusely rimmed with dark chromatophores;the belly is pale. The snout is darkened with a mix-ture of chromatophores and melanophores; thecheek is red with a slight yellow suffusion anteri-orly, two faint yellow spots below the anteroventralmargin of the eye; the ventral region of the cheekand the branchiostegal rays are a darker red. Thepectoral fin base has two reddish-orange spots, thedorsal of which lies a little anterior to the ventral,which latter is the darker of the two. The first dor-sal fin has a thin yellow-orange stripe just distal tothe base between the first three spines. The anteriormargin of a small (one-third pupil-diameter) ovoidblack spot overlies, or may begin just posterior to,the fourth dorsal spine and is confined to theregion between the fourth and fifth spines. It lieson the same plane as the stripe (Fig. 23), and has awide (about equal in width to the black spot) redhalo except dorsally over the spine, where the halogrades to yellow. A live specimen photographed at50 m is generally similar, but has a darker body, a

Fig. 24. Trimma papayum (live), Kawe Island, Raja Ampat. Photo by M. V. Erdmann.


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tral, and two spokes of a similar size and colour passdorsally towards the midline from the upper surfaceof the pupil.

C o l o u r p a t t e r n i n a l c o h o l : As above,but pale straw coloured (all red, orange and yellowareas pale), dark pigmentation outlining of scalepockets and in fin membranes remaining.Comparisons: Trimma papayum belongs to a

grade of 24 valid described species of Trimma, allof which have a Type B vertebral transition patternand scales present in the predorsal midline. In 11of these species, the second dorsal spine reaches nofurther posteriorly than the spine of the seconddorsal fin when the fin is adpressed (reaches poste-riorly to between the base of the fifth dorsal ray tobeyond the last ray in T. papayum). Four of theremaining species, T. lantana Winterbottom &Villa, 2003, T. mendelssohni (Goren, 1978), T. naudei Smith, 1957 and T. okinawae (Aoyagi,1949) have a deep postorbital trench (vs. a trenchabsent in T. papayum). Trimma randalli Winter-bottom & Zur, 2007, has an unbranched pectoralfin ray (vs. several middle rays branched) and T. flavicaudatum (Goren, 1982) and T. yanagitaiSuzuki & Senou, 2007, have 8 or fewer dorsal finrays (vs. 9). Of the six remaining species, T. barralliWinterbottom, 1995, has more lateral scales (25-26 vs. 23) and is confined to the Red Sea, T. imaiiSuzuki & Senou, 2009, has 16 pectoral fin rays (vs.18-19), T. nomurai Suzuki & Senou, 2007, has a

faint thin light stripe beneath the eye, light vermic-ulations over the snout and interorbital region, twopupil diameter orange spots on the opercle, and thered halo around the dark spot in the dorsal fin lacksthe yellow dorsal margin. A tiny black spot, situatedon the first dorsal spine just above its base, is presentin two of the 12 specimens, a 12.1 mm SL juvenileand an 18.9 mm SL female. A similar tiny black spotis present on and behind the second spine in a pho-tographed specimen from Maumere, Flores. Thesecond to sixth spines have elongate yellow spots alittle distal to the stripe and spot, and the fin mem-branes are invested with dark chromatophores andmelanophores, especially proximally and distally.The second dorsal fin has numerous yellow spots,the proximal (just above the base of the fin) seriesforming an irregular, interrupted stripe, the othersless obviously forming stripes, and the fin mem-branes contain numerous dark chromatophores andmelanophores with a reddish suffusion at the level ofthe proximal row of yellow spots. The anal and cau-dal fins are similar, with scattered yellow spots, a red-dish suffusion proximally, and numerous dark chro-matophores and melanophores (especially distally).The membranes of the pectoral and pelvic fins arehyaline, with the fin rays of the former reddish andthose of the pelvic fin yellowish. The dorsal three-quarters of the iris is dark red, with the ventral areabeing red and yellow. A thin charcoal to dark purpleline bisects the iris from anterodorsal to posteroven-

Fig. 25A-B. Head in A. left lateral (reversed) and B. dorsalview to show head papillae in Trimma xanthochrum (22.3mm SL male holotype, ROM 87483). Specimen stainedwith cyanine blue. Photo by R. Winterbottom.


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soft coral, 50 m, clove oil, 0900-0920, field #RW10-16, 28 January 2010, M. V. Erdmann.Paratypes: AMS I.45595-001 (formerly ROM85362), 4, 20.7-23.4, E. side of “Barracuda Rock”,200 m N of Wayil Island (02°11’43.4”S,130°25’37.9”E), 60 m, clove oil, 1300-1320, field# RW10-41, 1 February 2010, M. V. Erdmann.MZB 19781 (was ROM 85145), 25, 16.5-21.3,Jef Tsiep Island, west side, channel between it andsmall island to the west (00°23’05.7”S,130°16’37.1”E), 42 m, clove oil, 1600-1620, field# RW10-21, 20100128, M. V. Erdmann. ROM84885, 4, 7.4-25.1, Keruo Island off PenemuIsland, Fam Islands (00°35’15.4”S, 130°17’41.1”E), 70 m, clove oil, 0815-0850, field #RW10-04, 25 January 2010, M. V. Erdmann.ROM 84891, 2, 19.0-19.3, Keruo Island off Pen-emu Island, Fam Islands (00°35’15.6”S, 130°17’41.1”E), 56 m, clove oil, 1200-1300, field #RW10-05, 25 January 2010, M. V. Erdmann.ROM 85082, 8, 17.5-21.6, collected with theholotype. ROM 85132, 1, 16.6, Jef Tsiep Island,

pupil-diameter dark round spot on the bodybetween the pectoral and first dorsal fin (vs. such aspot absent), T. preclarum lacks scales on the oper-cle (vs. present) and has fewer anterior and poste-rior transverse scale rows (7-8 and 6-7 vs. 9-11 and8-9 respectively), and T. squamicana Winterbot-tom, 2004, has an unbranched fifth pelvic fin ray(vs. branched once) and lacks scales on the opercle(vs. present). Trimma halonevum Winterbottom,2000, is perhaps the most similar species to T.papayum in meristic values and in general shape,but adults have 2-3 cycloid scales on the cheek. Ithas a very similar small ocellated dark spot in thefirst dorsal fin, but this is confined to the mem-brane between the fifth and sixth spines (vs. on andjust posterior to the fourth spine), and has numer-ous similar dark haloed spots scattered over thebody (vs. such body spots absent). Distribution: Trimma papayum (Fig. 6) is cur-

rently known only from Maumere, Flores (photoonly) and Kawe Island in the Raja Ampat region ofIndonesia, where it was collected at a depth of 50 m.Etymology: Derived from the Spanish word

‘papaya’, a tropical tree whose fruit (papaya or paw-paw) has orange flesh and small black seeds, in allu-sion to the overall colouration and the small blackspot or spots in the first dorsal fin characteristic ofthe new species. Trimma papayum has been referredto informally (in litt.) as Trimma RW sp. 93.

Trimma xanthochrum n. sp.(Figs 2B, 6, 25-31)Step-spot pygmy goby


Trimma tevegae (Cohen & Davis, 1969) – Allen &Erdmann 2009: 619 (in part, Bird’s Head region,Indonesia)

Material Examined: A total of 13 collections, 123type specimens (7.4-25.1 mm SL), plus 19 non-types (including 7 tissue voucher specimens). Thedescription is based on up to 23 specimens fromROM 85082, 85155, 85208, 85225, 85319,87483 and 1834CS (17.7-24.1, 14 males, 7females, 2 unsexed).Holotype: ROM 87483, 22.3 mm SL male,Indonesia, Raja Ampat, about 1.2 km SSE of MutusIsland, on west side of sand spit (00°21’01.0”S,130°21’25.4”E), sponges, tunicates, some hard and

Fig. 26. Left lateral view of Trimma xanthochrum, 21.4 mm SL male paratype, ROM 85082, Mutus Island, Raja Ampat.Photo by R. Winterbottom.


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0750-0810, field # RW10-45, 2 February 2010, M.V. Erdmann. ROM 1834CS, 5, 9.9-20.9, collectedwith the holotype. USNM 399288 (formerly ROM85190), 3, 18.8-20.4, Tanjung Manare, WaigeoIsland, west side at about middle of width off smallcape (00°16’26.1”S, 130°19’01.5”E), sponges, hardcorals, 52 m, rotenone, 0750-0810, field # RW10-24, 29 January 2010, M. V. Erdmann.Additional (Non-type) Material. Seven specimens

for genetic analysis: one collected with ROM 84885(ROM T07708, 25.2); two with ROM 85082(ROM T07735, 20.7, T07736, 21.8), two withROM 85155 (ROM T07748, T07749, 21,8), onewith ROM 85208 (ROM T07752, 23.2) and onewith ROM 85319 (ROM T07761, 24.8). ROM84934, 12 (9.9-12.5), collected with the holotype).Diagnosis: Trimma xanthochrum is characterized

by a wide interorbital region (80-100% pupil diam-eter), a second dorsal spine reaching posteriorly tobetween the bases of the spine of the second dorsalfin and the fifth rays, 15-16 pectoral rays of whichthe middle 4-9 (usually 7-8) are branched, verticalrows of 2-3 sensory papillae below eye in rows 1-4and 4-5 papillae in row 5, a caudal spot which has aupper half about two-thirds the width of the lowerhalf, and usually an overall yellowish body with yel-low at least proximally in the caudal fin.Description: Dorsal fins VI + I 7-8 (once 7, n =

23), second and third spines longest, second reach-ing to between base of spine to fifth ray of seconddorsal fin (usually to between bases of second andthird rays), third reaching to between just anteriorto dorsal spine to base of first ray of second dorsalfin, rays all branched except posterior element oflast ray, last ray reaching to 35-45-50% distancebetween its base and first dorsal procurrent caudal

west side, channel between it and small island tothe west (00°23’05.7”S, 130°16’37.1”E), variety ofhard & soft corals, sponges, 19.8-22.9 m, clove oil,1600-1630, field # RW10-20, 28 January 2010, R.Winterbottom, L. Katz & P. Johannes. ROM85155, 13, 9.5-21.2, Tanjung Manare, WaigeoIsland, west side at about middle of width off asmall cape (00°16’26.1”S, 130°19’01.5”E), hard &soft corals, sponges, 19.8-22.9 m, rotenone, 0750-0820, field # RW10-23, 29 January 2010, R. Win-terbottom, L. Katz, P. Johannes, W. Kaka & W.Awom. ROM 85208, 2, 18.5-20.2, Wofoh Island,west coast near south end (00°15’21.9”S,130°17’32.0”E), many small corals, Tubastrea, seawhips, some black coral, sea fans, tunicates,sponges, some hydroids, 18.3-22.9 m, clove oil,1500-1540, field # RW10-25, 29 January 2010, R.Winterbottom & W. Kaka. ROM 85225, 21,15.2-22.0, Wofoh Island, west coast near southend (00°15’21.9”S, 130°17’32.0”E), many smallcorals, Tubastrea, sea whips, some black coral, seafans, tunicates, sponges, some hydroids, 12-16 m,clove oil, 1610-1700, field # RW10-27, 19 January2010, L. Katz. ROM 85319, 3, 22.3-24.1, Kepot-sol Island, north side at about middle of length, SEof Misool Island (02°09’32.1”S, 130°17’34.0”E),huge variety of soft & hard corals, some sponges,tunicates, & hydroids, 18.3-21.3 m, clove oil,0830-0900, field # RW10-37, 21 January 2010, R.Winterbottom, L. Katz & CI team. ROM 85333,17, 15.0-24.5, Kepotsol Island, east side(02°09’32.1S, 130°17’34.0”E), 66 m, clove oil,0830-0850, field # RW10-38, 1 February 2010, M.V. Erdmann. ROM 85389, 14, 16.7-25.0, SEislands off Misool, south side of Balbulol Island(02°01’29.5”S, 130°41’34.9”E), 45 m, rotenone,

Fig. 27A-B. Left lateral view of the caudal peduncle toshow differences in caudal spot of A. Trimma xanthochrum(20.9 female, ROM 85155) and B. T. tevegae (21.0 female,ROM 85320). Photo by R. Winterbottom.

Fig. 28. Left lateral view of Trimma xanthochrum, 20.8 mm SL male paratype, ROM 85155, Waigeo Island, Raja Ampat.Photo by R. Winterbottom.

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SD = 4.07, n = 20); pectoral fin 15-16 (x = 15.8,SD = 0.38, n = 23), 3-4-9 dorsal and 3-4-6 ventralunbranched rays respectively (more unbranchedrays in smaller specimens), middle rays branched,fin reaching posteriorly to a vertical line betweenposterior margin of anus and base of anal spine;pelvic fin I 5, no frenum, basal membrane 8-17%length of fourth ray (x =12.3%, SD = 2.32, n =17)), first four rays with one sequential branch,fifth ray usually branched once dichotomously(unbranched in 3 of 23) and 45-58% length offourth (x = 51.1%, SD = 3.78. n = 18), fourth rayreaching posteriorly to between bases of anal spineto fourth anal-fin ray. Lateral scales 23-24 (x =23.2, SD = 0.36, n = 20), when 24, first scale inpectoral fin axil is half the size of next scale andcycloid; anterior transverse scales 9; posterior trans-verse scales 8; 12-14 irregular scale rows in midlineof predorsal (x = 12.8, SD = 0.54, n = 20); threerows of cycloid scales on cheek, dorsalmost row of3-4, middle row of 8-9 and ventralmost row of 3-6 scales (usually 3, 8 and 5 respectively – Fig. 25A);usually four irregular horizontal rows of scales onopercle, with 1-3-4, 3-4, 2-4 and 1-2 cycloid, mid-dle scales of second row may be ctenoid, once afifth row of a single cycloid scale; 3 vertical rows ofcycloid scales on pectoral base with 4 scales (once3) in posteriormost row; 7-8-9 (x = 7.5, SD = 0.59,n = 20) cycloid prepelvic scales; 11-12 (x = 11.8,SD = 0.38, n = 17) circumpeduncular scales; 9-10scales in midline between base of last anal ray andfirst ventral procurrent ray (x = 9.9, SD = 0.31, n= 19); body scales ctenoid except for cycloid scaleson anterior belly midline, beneath and just poste-rior to pectoral-fin base, and one or two scales adja-cent to posterodorsal margin of orbit. Gill opening

ray (x = 42.0%, SD = 4.19, n = 20); anal fin I 8,first ray and posterior element of last ray in 17 (of23) unbranched; posteriormost ray reachingbetween 28-35-43% distance between its base andfirst ventral procurrent caudal ray (x = 35.4%,

Fig. 30. Left lateral view of Trimma xanthochrum, 23.2 mm SL male paratype, ROM 85319, Kepotsil Island, Raja Ampat.Photo by R. Winterbottom.

Fig. 29. Left lateral view of Trimma xanthochrum, 22.0 mm SL female paratype, ROM 84885, Penemu Island, Raja Ampat.Photo by R. Winterbottom.

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anteroventral and posteroventral margins of eye.Papillae immediately below eye on cheek in fivetransverse rows, usually with 2, 2, 3, 3 and 5 papil-lae in vertical rows 1 through 5 (Fig. 25A – see alsoFig. 2B). Tongue truncate with rounded edges anda shallow V-shaped tip. Gill rakers on first arch 3-4 + 14-17 = 18-21 (x = 3.9 + 15.5 = 19.4, SD =0.28, 0.72, 0.87 respectively; n = 23). Anteriornasal opening a broad tube extending almost toabove anterior margin of upper lip, posterior nasalopening a pore with a raised rim, both protrudingfrom slightly raised oval sac, nasal apparatus con-fined to anterior one-third of snout. Bony interor-bital 80-89-100% pupil width (mean = 91.4%, SD= 6.67, n = 19), interorbital shallowly concave witha broad, rounded median fleshy ridge and no pos-torbital trough or trench (Fig. 25B). No dermalcrest anterior to first dorsal fin. Epaxialis muscula-ture extending anteriorly to above posterior marginof pupil in dorsal midline. Vertebral transition a

extending anteroventrally to a vertical below poste-rior one-third of pupil. Upper jaw teeth of a row ofenlarged spaced, curved caniform teeth thatdecrease about 50% in size from symphysis to dis-tal tip of premaxilla, two to three inner rows ofsmaller conical teeth at symphysis, innermost rowslightly larger and curved, grading to a single rowposteriorly to end at distal tip of premaxilla. Lowerjaw teeth with an outer row of 6-8 enlarged curved,spaced canines ending at bend of dentary, aboutfour irregular rows of small conical teeth at symph-ysis, innermost row increasing in size posteriorlyuntil almost as big as outer symphysial row at mid-dentary, then gradually decreasing in size and end-ing at dorsal tip of coronoid process. Sensory papil-lae very delicate and subject to abrasion, especiallythose on cheek, opercle and immediately aboveopercle and posterior to eye, several specimens withsupernumary papillae (not included in counts inTable I) which were most frequently found around

Fig. 31. Trimma xanthochrum (live) Triton Bay, Papua. Note the second spine of the first dorsal fin is deformed at its tip.Photo by G. R. Allen.


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where the entire scale pockets are heavily outlinedwith dark pigment. A diffuse dark blotch coversthe hypural region and bases of the caudal rays anda thin, diffuse vertical yellow bar over the ends ofthe hypurals separates the blotch into anterior andposterior halves, the posterior half being morestrongly pigmented than the anterior half. Thedorsal half of the blotch is about two-thirds thewidth of the ventral half (Fig. 27A), and does notextend anteriorly beyond the base of the anterior-most procurrent fin ray. The first dorsal fin has aone-third pupil width dark basal stripe, followedby a pupil width yellow stripe, with the rest of thefin heavily invested with melanophores; the spinesare reddish. The second dorsal fin has a similardark basal stripe, above which a narrow yellowstripe expands posteriorly until it occupies theentire height of the fin from the sixth fin ray on;the distal anterior portion of the fin has numerousmela no phores. The anal fin is essentially hyalinewith a slight reddish suffusion on the posterior-most rays. The pectoral fin is yellow to orange witha few scattered brown chromatophores except onthe dorsal rim, which is rimmed with melano -phores to form a diffuse, elongate spot; the fin ishyaline with red fin rays. The pelvic fin is dusky.The dorsal and ventral margins of the caudal finare hyaline with reddish unbranched caudal rays,

modified Type A, with the tenth abdominal andfirst caudal vertebrae (only) with a small basalhaemal canal, the main haemal arch of the firstcaudal vertebra expanded, that of the second abouthalf the size of the first, remaining caudal haemalarches normal; swimbladder terminates at thehaemal canal of second caudal vertebra.

C o l o u r p a t t e r n (based on freshly collectedmaterial, a total of seven images, 4 males, 3females, and one live specimen). A 21.4 mm SLmale (Fig. 26) has yellowish body (especially alongthe ventral region) with a diffuse rosy stripe abovethe lateral septum to within about a scale width ofthe dorsum. The lining of the swim bladder isdarkly pigmented, which shows through the bodywall and is augmented by brown subdermal chro-matophores over the upper half of the abdominalcavity, together forming a wedge-shaped darkerstripe which fade out just posterior to the anal fin.The upper part of the head and upper jaw is heav-ily and darkly pigmented, with a thin diffuse redstripe just below the eye from the vertical limb ofthe preopercle to middle of the maxilla. The areabelow this stripe is light red with yellow suffusions,becoming pale ventrally. The body scales are thinlyedged with melanophores along their exposed mar-gins, except ventrally on the abdomen and trunk,where there is no such pigmentation, and dorsally,

the rest of the fin is yellow with a pale central prox-imal stripe and the membrane between the middletwo rays has a red suffusion. The iris is densely mot-tled with dark pigment, with a golden suffusion pos-teroventral to the pupil; a dark stripe passes over thetop of the pupil, above which is a diffuse reddish-yellow stripe. A 20.8 mm SL male (Fig. 28) differsin having the dorsal half of body pale, the thin redstripe below the eye is followed by a diffuse palestripe just below it, and the lower jaw, except the tip,is pale. The dorsal half of the caudal blotch is betterdeveloped and about four-fifths the width of theventral half; the proximal half of the caudal fin ismuch more intensely pigmented than the distal half,and the central light stripe is better developed andreaches almost to the distal margin. The iris is mot-tled with black, red and orange pigment with a veryclear light stripe margining the dorsal edge of thepupil, above which the iris is dark red. A 22.0 mmSL female (Fig. 29) is much yellower overall, withthe intense yellow pigment in the caudal fin extend-ing almost to the distal margin of the fin and a sprin-kling of melanophores in the central light stripe.The caudal blotch is somewhat better defined, anddisparity in width between the upper and lowerhalves more obvious. The iris is a mix of golden andblack pigment, and the stripe over the pupil is black.A 23.2 mm SL male (Fig. 30) has an overall reddishcast, grading to orange-red below the dark wedge-shaped internal stripe, and the yellow on the caudalfin is confined to the proximal half of the fin.Among the remaining photographed specimens,essentially the same variation is apparent, althoughthe light stripe under the eye may have a bluish cast.A specimen photographed live underwater at TritonBay (Fig. 31) is overall reddish-orange, with diffusebody stripes (a somewhat darker mid-lateral stripewith light orange-yellow dorsal and ventral bodystripes), an oblique white stripe across the iris overthe top of the pupil, and greyish margins on theunbranched rays and a posteriorly attenuating greystripe in the center of a yellow caudal fin.

C o l o u r p a t t e r n i n a l c o h o l : As above,but pale straw coloured with only the darkly pig-mented areas remaining visible. The dark pigmenta-tion on the dorsal surface and the caudal spot areespecially evident.Comparisons: This new species is a member of the

T. tevegae species group, defined by an interorbitalwidth of greater than 75% of the pupil width and aType A abdominal/caudal vertebral transition.Among the described species in this group, it differs

from all but T. caudomaculatum and T. tevegae inhaving a well developed large dark spot at the end ofthe caudal peduncle that extends the full width ofthe peduncle. In both those species, the caudal spotis rounded with both dorsal and ventral halvesapproximately equal in diameter (Fig. 27B – vs. dor-sal half about two-thirds the width of the ventralhalf – Fig. 27A), there are 12-14 unbranched pec-toral fin rays (vs. 15-16 rays with at least 4, and usu-ally 6-8, of the middle rays branched), the fifthpelvic ray is nearly always unbranched (vs. branchedonce in 87% of specimens), and the vertical papillaein rows 2-4 just below the eye consist of a singlepapilla each (vs. 2-3 papillae in each row, see Fig. 2B and 26A). In T. caudomaculatum, the second spineof the first dorsal fin extends much further posteri-orly, reaching to the caudal peduncle. In contrast,the second spine of T. tevegae is not elongate, reach-ing posteriorly only as far as the spine of the seconddorsal fin when the fin is adpressed. An analysis ofthe CO1 gene of 72 specimens originally identifiedas T. tevegae from three localities (Raja Ampat, Palauand Japan) suggests that there are seven deeply diver-gent haplogroup clusters, separated from each otherby between 9.1-22.9% of the standard mitochondr-ial 5’ COI barcode locus (Winterbottom et al., inprep.). One of these groups, informally called“Group 2”, is the species described here as T. xan-thochrum, and it is separated from all the others by13.8-17.8%. It is impossible to say at presentwhether the true T. tevegae (type locality RabaulHarbour, New Britain) is represented in the samplesanalysed, since no specimens from that locality arecurrently available for genetic analysis. The collec-tions analysed to date from both Palau and fromRaja Ampat contain three deeply divergent hap-logroups, each characterized by extremely lowwithin-cluster variation (i. e. < 1%). It is thereforepossible or even probable that more than one ofthese (or other) clusters may be present at the typelocality of T. tevegae, but if so, it is unknown at thispoint which one, if any, might represent the typespecies. Morphological or colour differences betweenthese groups, if they exist, have not been established,except for T. xanthochrum and the apparent differ-ence in the degree of development of the second dor-sal spine between T. caudomaculatum and T. tevegae.Distribution: Trimma xanthochrum is currently

only positively recorded from the Bird’s Head regionof Indonesia (Fig. 6) in depths of 12-70 m.Etymology: The name is derived from the Greek

‘xanthos, yellow or golden, and ‘chros’ colour of the

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skin, or body surface, in allusion to the yellowcolouration of the body and caudal fin. This specieshas been referred to informally (in litt.) as TrimmaRW sp. 94.

ACKNOWLEDGEMENTSI would especially like to thank Mark V. Erdmann

(Conservation International’s Indonesia MarineProgram) for the invitation to participate in thefieldwork, and for diligently collecting additionalmaterial after the trip in which I participated. Iwould also like to express my gratitude to LaureKatz who, in addition to collecting many specimensof gobies for me, made sure that I always returnedsafely to the surface after each dive. My sincerestthanks to the other members of the “fish” team(Rudi Dimara, Andi Fauzan, Christine Huffard,Muhammad Lazuardi and Defy Pada), to the cap-tain and crew of the Putri Papua for their unstintinghelp, support and companionship during the survey,to the Indonesian Department of Nature Conserva-tion (PHKA), the Indonesian Institute of Sciences(LIPI), and to the Raja Ampat government for theirsupport of this work. I also extend my gratitude tothe Paine Family Trust for their financial support ofthis survey through Conservation International’sIndonesia Marine Program. Thanks to Doug Hoeseand Peter Miller for their comments on the sectionon papillae in Trimma, and to Gerry R. Allen andMark V. Erdmann for permission to use their excel-lent photographs. It is always a pleasure to acknowl-edge the tremendous contributions of Margaret Zur(ROM), who for many years gathered and analyseddata from the huge collections of borrowed andROM Trimma. Her notes have been invaluable inmaking the comparisons of the new species withpreviously described members of the genus. Addi-tional financial support for this study was generouslyprovided the ROM’s Department of Natural His-tory and my NSERC Discovery Grant A7619.

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AHNELT, H. & GÖSCHEL, J. 2003. Morphological differencesbetween the eastern Pacific gobiid fishes Quietula guay-masiae and Quietula y-cauda (Teleostei: Gobiidae) withemphasis on the topography of the lateral-line system.Cybium 27 (3): 185-197.

AKIHITO, P., HAYASHI , M. & YOSHINO, T. 1988. SuborderGobioidei. In: The Fishes of the Japanese Archipelago. (Eds.

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AKIHITO, SAKAMOTU, K., IWATA, A., AND IKEDA, Y. 1993.Cephalic sensory organs of the gobioid fishes. In: Fishesof Japan with pictorial keys to the species. (Ed. T. Nakabo).Tokai Univ. Press, Tokyo, Japan [In Japanese], pp 1088-1116.

AKIHITO, SAKAMOTO, K., IKEDA, Y., & SUGIYAMA, K. 2002.Gobioidei. In: Fishes of Japan with pictorial keys to the species.(Ed. T. Nakabo). English edition, Vol. II, Tokai UniversityPress, Tokyo, pp 1139-1310; 1596-1619.

ALLEN, G. R. & ERDMANN, M. V. 2009. Reef fishes of theBird’s Head Peninsula, West Papua, Indonesia. Check List5 (3): 587-628.

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AURICH, H.J. 1938. Mitteilung XXVIII der Wallacea-Expe-dition Woltereck. Die Gobiiden. (Ordnung : Gobioidea).Internationale Revue der gesamten Hydrobiologie und Hydro-graphie, 38(1/2) :125-183.

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DIMARA, R., FAUZAN, A., LAZUARDI, M., PADA, D., ALLEN,G. R., ERDMANN, M. V., HUFFARD, C. L., KATZ, L. S. &WINTERBOTTOM, R. 2010. An illustrated list of additionsto the goby fauna of the Raja Ampat Islands, Indonesia(Pisces, Gobiidae). Check List 6 (4): 619-625.

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WINTERBOTTOM, R. 2000. Four new species of Trimma(Gobiidae), from the Indian and western Pacific oceans.aqua, Journal of Ichthyology and Aquatic Biology 4 (2): 57-66.

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WINTERBOTTOM, R. & BURRIDGE. M. 1992. A revision ofEgglestonichthys and the species of Priolepis possessing atransverse cheek papillae pattern (Teleostei; Gobiidae); witha discussion of relationships. Canadian Journal of Zoology70: 1934-1946.

WINTERBOTTOM, R. & Villa, C. A. 2003. A new species ofthe Trimma caesiura complex (Gobiidae, Teleostei) fromthe north-eastern margin of the Australian Plate, with aredescription of the other nominal species in the com-plex. aqua, Journal of Ichthyology and Aquatic Biology 7(1): 13-28.

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KIMURA, S. & MATSUURA, K. 2003. Fishes of Bitung, North-ern Tip of Sulawesi, Indonesia. Ocean Research Institute,University of Tokyo. vi + 244 pp.

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KUITER, R. H. & TONOZUKA, T. 2004. Pictorial Guide to:Indonesian Reef Fishes. Part 3. PT Dive & Dive’s, Bali,Indonesia. iv + 623-893.

MILLER, P. J. 1972a. Generic status and redescription of theMediterranean fish Gobius lichtensteini Kolombatovic,1891 (Teleostei: Gobioidea), and its affinities with certainAmerican and Indo-Pacific gobies. Journal of Natural His-tory 6: 395-407.

MILLER, P. J. 1972b. The identity of Gobius affinis Kolom-batovic, with notes on the systematics and biology ofPomatoschistus pictus (Malm) (Pices: Gobiidae). Annali DelMuseo Di Storia Naturale Di Genova 79:53-88.

MILLER, P. J. 1973 The identity of Gobius affinis Kolombat-ovic, with notes on the systematics and biology ofPomatoschistus pictus (Malm) (Pisces: Gobiidae). Annali delMuseo Civico di Storia Naturale `Giacomo Doria’ 79 (for 1Sept 1972): 53-88.

MILLER, P. J. & WONGRAT, P. 1979. A new goby (Teleotei:Gobiidae) from the South China Sea and its significance forgobioid classification. Zoological Journal of the Linnean Soci-ety 67: 239-257.

SANZO, L. 1911. Distribuzioine delle papille cutanee (organiciatiformi) e suo valore sistematico nei Gobi. Mitteilungenaus der Zoologische Station zu Neapel 20 (2): 251-328.

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SUZUKI, T. & Senou, H. 2007. Two new species of the gob-iid fish genus Trimma (Perciformes: Gobioidei) from south-ern Japan. Bulletin of the National Museum of Natural Sci-ence, Ser. A, Suppl. 1: 175–184.

SUZUKI, T. & SENOU, H. 2008. Two new species of the gob-iid fish genus Trimma (Perciformes: Gobioidei) from south-ern Japan. Bulletin of the National Museum of Natural Sci-ence, Ser. A, Suppl. 2: 97-106.

SUZUKI, T. & SENOU, H. 2009.Two new species of the genusTrimma (Perciformes: Gobioidei) from Japan and Palau.Bulletin of the Kanagawa Prefectual Museum (Natural Sci-ence) (38): 107-112.

TAKAGI, K. 1988. Cephalic sensory canal system of the gob-ioid fishes of Japan: comparative morphology with special


A redescription of the gobiid fish Cryptocentrus sericus Herre, with clarificationof Cryptocentrus leptocephalus and C. melanopus

Doug Hoese1, Koichi Shibukawa2 and Jiro Sakaue3

1) Australian Museum, 6 College Street, Sydney, NSW 2010, Australia. Email: [email protected]

2) Nagao Natural Environment Foundation, 3-10-10 Shitaya, Taito-ku, Tokyo 110-0004, Japan.E-mail: [email protected]

3) Southern Marine Laboratory, P.O. Box 1598 Koror, Republic of Palau 96940. E-mail: [email protected]

Received: 12 March 2011 – Accepted: 13 May 2011

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d’autres espèces aux données morphologiques analogues.Des illustrations de C. sericus sont fournies ainsi que de l’espèce la plus voisine, C. fasciatus. Une brève comparai-son et des dessins sont prévus pour séparer les espècesproches C. leptocephalus et C. melanopus. Ces deux espècesont été confondues et souvent identifiées erronément.

SommarioCryptocentrus sericus Herre è ridescritto utilizzando

l’olotipo e materiale rinvenuto recentemente nelle col le -zioni di Australian Museum, Museum and Art Gallery ofthe Northern Territory e Western Australian Museum. Laspecie si distingue per la colorazione, la presenza di scagliepredorsali e per il numero di scaglie e di raggi delle pinne.Si fornisce una comparazione con altre specie aventi mor-fologia simile. Sono mostrate immagini di C. sericus e dellaspecie più affine, C. fasciatus. Sono altresì fornite una brevedescrizione comparativa e alcune figure che aiutano a di -stinguere altre due specie molto vicine, C. leptocephalus eC. melanopus, spesso erroneamente identificate e confusetra loro.

INTRODUCTIONCryptocentrus contains 34 described and recog-

nised species. A few species, which live aroundcoral reefs, are generally well known, but most ofthe species live in silty environments adjacent toreefs, mangroves or on the continental shelf. Manyare known only from a few trawled specimens.Species found in shallow waters are known to livewith alpheid shrimps. Information is presentedhere to clarify the identity of some of the previ-ously described species, which have been confusedor believed incorrectly to be undescribed. Three ofthe species treated here live adjacent to coral reefs,but often in silty waters well away from coral. One

AbstractCryptocentrus sericus Herre is redescribed from the holo-

type and recently collected material in the AustralianMuseum, Museum and Art Gallery of the Northern Terri-tory and Western Australian Museum. The species is dis-tinctive in coloration, presence of predorsal scales, and infin-ray and scale counts. We provide a comparison withother species with similar morphological features. Illustra-tions are provided of C. sericus and its most similar species,C. fasciatus. A brief comparison and figures are providedfor separating the related C. leptocephalus and C. melano-pus. These two species have been confused and frequentlymisidentified.

ZusammenfassungDie Beschreibung von Cryptocentrus sericus Herre wird

nach dem Holotypus und nach neuerem Material im Aus-tralischen Museum, in Museum und Kunstgalerie desNördlichen Territoriums und im WestaustralischenMuseum revidiert. Die Art unterscheidet sich durch Farb -gebung, Vorhandensein prädorsaler Schuppen, Flossen-strahlen und Schuppenzahlen. Der Vergleich mit anderenArten mit ähnlichen morphologischen Kennzeichen wirdermöglicht. Abbildungen zeigen C. sericus und die ammeisten ähnliche Art C. fasciatus. Durch eine kurzeErläuterung und weitere Zahlen wird die Abgrenzung zuden nahe verwandten C. leptocephalus und C. melanopusermöglicht. Die beiden Arten wurden oft vermischt undfalsch bestimmt.

RésuméCryptocentrus sericus Herre est redécrit sur base de l’holo-

type et de matériel récemment réuni provenant de l’Aus-tralian Museum, du Museum and Art Gallery of theNorthern Territory et du Western Australian Museum. L’espèce se distingue par la coloration, la présence d’écaillesprédorsales et le nombre de rayons des nageoires et d’écailles. Nous avons prévu une comparaison avec

A redescription of the gobiid fish Cryptocentrus sericus Herre, with clarification of Cryptocentrus leptocephalus and C. melanopus

164aqua vol. 17 no. 3 - 10 July 2011

Cryptocentrus sericus Herre, 1932Figs 1-5

Cryptocentrus sericus Herre, 1932: 440 (type locality:Canton market, China).

Cryptocentrus sp. 1 – Akihito in Masuda et al. 1984:259, pl 243L (Japan, hoshizorahaze).

Cryptocentrus sp. 3 – Okamura & Amaoka 1997:608 (Japan).

Cryptocentrus sp. – Hayashi et al. 1990: 135, pl. 2,fig. 4 (Amami-oshima Island, Ryukyu Islands,Japan and introduced new Japanese name of Futa-hoshi-takanoha haze); Hayashi & Shiratori 2003:133 (Japan, ventral barred goby); Kuiter & Debe-lius 2006: 653 (Indonesia, Ventral-Barred Goby).

Cryptocentrus sp. B. – Myers 1999: 240, pl.152G, H(Yaeyamas and Palau); Suzuki et al. 2004: 322(Japan).

Cryptocentrus n.sp. 1 – Larson, Jafaar & Lim 2008:148 (Singapore).

Material Examined: HOLOTYPE: CAS-SU25725, 75.5 mm SL female from fish market atCanton, China. OTHER MATERIAL. HongKong, China NTM (uncatalogued) 2(66–70):Australia: AMS I.19474-003, 1(39), Linnett Reef,Queensland, 25 m; WAM P.31650-006, 3(23-67),Cassini Island, Western Australia, 20-30 m. Philip-pines: WAM P.32882-007, 1(55), Palawan, PinasilIsland, 15-17 m; WAM P.33000-003, 1(58),Palawan, Imorigue Channel, 12-15 m. Indonesia:WAM P.31303-004, 1(43), Bintan Island,Carston’s reef, 12 m; WAM P.31525-007, 1 (68),Sumatra, 15-40 m; WAM P.32814-002, 1(38),Papua, Pulau Batuputih, Fak Fak coast, 3-5 m.Brunei: WAM P.33030-003, 2(35-42), Bruneipatches east, 12-15 m. Thailand: AMS I.42904-003, 1(71), Ka Island, east coast; AMS I.42911-001, 3(54-79), Pu Island, northwest coast; AMSI.42911-002, 2(39-40). In addition, 41 distribution records of this species

can be found under the name Cryptocentrus sp. 2at FishPix of the Kanagawa Prefectural Museum ofNatural History, Odawara and the Museum of Sci-ence and Nature, Tokyo (http://fishpix.kahaku.go.jp/fishimage-e/search.html), as follows:Amami-oshima Island, Ryukyu Islands, Japan:KPM-NR 36538; KPM-NR 40704. Ishigaki-jimaIsland, Yaeyama Group, Ryukyu Islands, Japan:KPM-NR 81680 (yellow phase). Iriomote-jimaIsland, Yaeyama Group, Ryukyu Islands, Japan:KPM-NR 81680. Okinawa Island, Ryukyu

of these species is well known and often pho-tographed, and typically is referred to as the ventralbarred goby. Workers have suspected that it repre-sented an undescribed species, but evidence is pre-sented here that it was described over 75 years ago.Cryptocentrus is superficially similar to Ambly -

eleotris, and the two genera are often difficult toseparate in the field. Hoese and Steene (1978) pro-vided diagnoses of the two genera and noted thatCryptocentrus has the upper longitudinal cheekpapilla row extending forward to the second orthird transverse row, under the eye (vs. under fifthor sixth row); the mandibular papillae are arrangedin two parallel rows extending posteriorly on thesides of the chin (vs. a single papillae set in a pit ateach side of chin); the gill opening is morerestricted, usually below or in front of the posteriorpreopercular margin (vs. behind the margin); scalesare normally cycloid, except for 2 or 3 species (vs.always with some ctenoid scales); dorsal and analrays vary from I,9 to 12 (vs. I,12-19). Randall(2004) incorrectly reported the papillae set in pits,first reported by Hoese & Steene (1978) on thechin, as being behind the upper jaw. He alsoreported it as unique, but the same pattern foundin Amblyeleotris occurs in Ctenogobiops, as illus-trated by Randall, Shao & Chen (2003) and Van-derhorstia, as illustrated by Shibukawa & Suzuki(2004). Hoese and Larson (2004) provided addi-tional diagnostic features for Cryptocentrus. Crypto-centrus species generally have a larger mouth reach-ing well behind the eyes (versus usually endingbelow eye) and most Cryptocentrus species aredeeper bodied and have a proportionally largerhead, than those of Amblyeleotris. In Cryptocentrusthe lower rakers on the inner face of the first archand outer face of the second arch are not ossified,but in Amblyeleotris they are ossified. Unossifiedrakers are rare in gobioid fishes, but are also knownin Callogobius.

METHODSMethods of counts and measurements follow

Hoese & Larson (2004). Institution abbreviationsfor material examined follows Leviton et al.(1985). After the institution catalogue number thenumber of specimens is given followed by the sizerange in mm SL (standard length). Counts for theholotype are marked with an asterisk. All measure-ments were taken with dial callipers and expressedas range of percent standard lengths.

Fig. 1. Freshly collected specimen of Cryptocentrus sericus, AMS I.19474-003; note that as the specimen was speared theimage has been modified and the belly and upper part of the pelvic fin is distorted.

Fig. 2. Holotype of Cryptocentrus sericus, CAS SU 25725, 75.5 mm SL female.

Doug Hoese, Koichi Shibukawa and Jiro Sakaue

165 aqua vol. 17 no. 3 - 10 July 2011

17m. Sabah, Borneo: KPM-NR 27873, 1m;KPM-NR 31690, 22 m.Diagnosis. Mental frenum absent. Mouth large;

ending well beyond end of eye (by about one pupildiameter or slightly less); jaws forming angle of 22-28° with body axis; upper margin of upper jaw inline with point about one-quarter to one thirdpupil diameter below eye. Cheeks slightly bulbous.Fleshy interorbital very narrow, much less than halfpupil diameter. Gill opening reaching to below apoint just under posterior preopercular margin.Head papillae minute and not on distinct ridges.Scales entirely cycloid. Predorsal area partly scaled,with scales reaching forward to just before poste-rior preopercular margin to midway between eyeand posterior preopercular margin. Cheek nakedor with one or two minute embedded cycloidscales; operculum naked or with 1-5 small embed-ded cycloid scales. Pectoral fin base usually naked,sometimes with small patch of scales centrally. Pre-pelvic area partly to fully scaled with 6-16 rows

Islands, Japan: KPM-NR 7112, 25 m; KPM-NR11688, 20m (yellow phase); KPM-NR 15880,20m (yellow phase); KPM-NR 28237, 25m;KPM-NR 52478; KPM-NR 61359 (yellowphase); KPM-NR 69230, 3 m; KPM-NR 80827, 7m. Zamami-jima Island (Ke ra ma Group), RyukyuIslands, Japan: KPM-NR 42170; KPM-NR87476, 17 m. Hong Kong: KPM-NR 34758;KPM-NR 34759; KPM-NR 34760 (yellowphase); KPM-NR 34761 (yellow phase); KPM-NR34762; KPM-NR 34763; KPM-NR 64667; KPM-NR 64669; KPM-NR 64712, (yellow phase);KPM-NR 69167, 3m; KPM-NR 69178, 2m;KPM-NR 69187, 4m; KPM-NR 69191, 4m;KPM-NR 69192, 4m, (yellow phase); bKPM-NR69196, 4m. Negros Island, Philippines: KPM-NR35855 (yellow phase); KPM-NR 35856 (yellowphase); KPM-NR 35857; Boracay Island, Philip-pines: KPM-NR 61019 (yellow phase); KPM-NR80440, 15m (yellow phase). Sulawesi, Indonesia:KPM-NR 11476, 15-20m; KPM-NR 85968, 6-

Fig. 3. Underwater photo of Crytocentrus sericus from the Great Barrier Reef. Photo by R. Steene.


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groove behind, about twice snout length. Anteriornostril at end of short tube, with broad flat rim,just above upper margin of upper lip. Posteriornostril a large pore about one nostril diameterbehind anterior nostril, and one nostril diameterfrom eye. Preoperculum long, distance from end ofeye to upper posterior preopercular subequal to dis-tance from snout to an eye length behind eye. Pos-torbital long, slightly shorter than distance from tipof snout to posterior preopercular margin. Bodyslender, depth at anal origin 16.7-18.1% SL. Upperjaw long, reaching about one pupil diameter behindeye (13.3-16.1% SL). Teeth conical and curved.Teeth in upper jaw: outer row of teeth curved,enlarged and wide-set, teeth larger near angle of jaw;three to four rows of smaller, depressible teeth ante-riorly, tapering to two to three rows posteriorly, teethin rows pointing inward into mouth; an innermostrow of one to two enlarged backwardly directedteeth anteriorly on each side of jaw. Teeth in lowerjaw: teeth in outer row curved, conical, wide-set,covering anterior end of dentary only, three to fourinner rows of smaller conical teeth anteriorly andtwo to three rows posteriorly, innermost row of teethenlarged and larger than teeth in outer row, withlargest teeth just behind bend in dentary. Tongue tiprounded. Gill rakers on outer face of first arch slen-der, denticulate on posterior margin, rakers muchshorter than filament length. Rakers on inner face offirst arch and other arches short and denticulate atdistal tip. Body covered with cycloid scales. Headlargely naked, midline of nape with scales extendingto above preoperculum, operculum and preopercu-lum naked or with a few small embedded scales.Adpressed length of first dorsal fin 22.2-28.9% SL,

before pelvic fin origin. Belly fully covered withcycloid scales, except for small naked area justbehind pelvic insertion. First dorsal fin moderatelyhigh, with truncate margin, first spine longest butnot distinctly prolonged into filament. Cheek oftenwith two short black stripes, one just above posteriorend of jaws and second just above lower stripe. Firstdorsal fin with two black, oblique stripes. Pelvic finwith 4-6 curved black bars in females, fin usuallyuniformly dark in males or with one or two darksbands posteriorly. Pelvic fin large; reaching to justshort of anus. Second dorsal-fin rays usually I,10;anal fin rays usually I,9; pectoral fin rays 16-17; lon-gitudinal scale count 68-74; predorsal scale count20-27; transverse scale count (TRB) 21-27.Description: Based on 16 specimens, 28-79 mm

SL. Counts of holotype indicated by asterisk.Numbers in parentheses after counts indicate thenumber of specimens with that count.First dorsal spines VI(16)*; second dorsal rays

I,9(1), I,10(14)*, I,11(1); anal rays I,9(16)*; pec-toral rays 16(3), 17(13)*; longitudinal scale count64(2), 65(2), 67(1), 68(1), 70(1)*, 71(1), 74(1),78(1); predorsal scale count 18(1), 19(1)*,20(3),22 (4), 23(1), 24(1); transverse scale count (TRB)21,(1), 24(2), 25(2), 26(4)*, 27(1); gill rakers onouter face of first arch 2+10(1), 3+1+10(1),2+1+11(1), 2+1+12(1), 4+1+12(1)*; gill rakers onouter face of second arch 3+1+12(1), 3+1+13(1),4+1+13(1), 3+1+15(1)*; segmented caudal rays9/7(1) (2), 9/8(13)*; branched caudal rays7/6(12)*, 7/7(2); vertebrae 10+16 (holotype).Head strongly compressed. Snout rounded in

dorsal view; steeply oblique (slightly convex) inside view. Eye large and elevated, with shallow

Fig. 5. Underwater photo of yellow phase of Cryptocentrus sericus from Sulawesi. Photo by R. Steene.

Fig. 4. Sensory papillae of head of Cryptocentrus sericusbased largely on holotype.


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middle of eye; middle pore midway between upperand lower pores.Papillae, based largely on holotype. Head papilla

pattern transverse (Fig. 4). Cheek with five VT lines;first from below middle of eye to middle of jaws; sec-ond incomplete and extending upward from justbehind first row, not meeting eye, ending just belowor before upper LT line; third interrupted by upperLT line; fourth slightly oblique cut into two parts byupper LT line with upper part nor meeting upper LTline; fifth a short oblique line before infraorbital poreand a vertical section ventrally below upper LT line.Upper LT reaching to near posterior preopercularmargin, lower LT line ending under or just behindfifth VT line. An oblique line extending above upperjaw toward anterior nostril. Three short oblique linesfrom anterior part of upper jaw to first VT line. Pre-opercular-mandibular series with outer LT line inter-rupted just behind upper jaw; inner line composed ofsingle row of papillae, not interrupted behind jaws. Atransverse (TT) line behind each eye. Chin papillaearranged in two posteriorly converging LT lines.C o l o r a t i o n i n a l c o h o l . Head and body

light brown. Head with two dark brown to black,short, horizontal stripes above the posterior two-thirds of the upper lip, faint or absent in specimensabove 55 mm SL; a dark brown oval spot abouthalf pupil diameter at posterior end of preopercu-lum, followed by similar sized spot anteriorly onoperculum, spots often fused to form larger ovalspot or short stripe, often faint or absent in speci-mens over 55 mm SL; dark brown oval spot, ori-ented obliquely on upper operculum; dark brownto black stripe above; sometimes with oval darkspot, height about half pupil diameter behind eye;

origin just above pelvic fin insertion. Second dorsalfin base 29.3-32.8% SL. Anal fin base 20.0-23.8%SL. Pectoral fin with rounded margin, reaching toabove or just before pelvic fin tip, to or just beforeanus in adults, length, 22.2-28.0% SL. Pelvic disclarge, reaching to anus or slightly beyond, but not toanal fin origin; pelvic length 23.3-29.0% SL inadult. Caudal fin with rounded posterior margin,central rays longest; length 26.7-32.7% SL.Head pores. Posterior nasal pore slightly median to

posterior nostril; median anterior interorbital porejust before eyes; median posterior interorbital aboveposterior margin of pupil; postorbital pore behindupper quarter of eye; infraorbital pore below andbehind postorbital pore, behind and in line withupper margin of pupil; lateral canal pore above mid-dle of preoperculum; terminal lateral canal poreabove posterior preopercular margin; a long tubewith pores at each end above operculum; three pre-opercular pores, upper in line with point just below


168aqua vol. 17 no. 3 - 10 July 2011

lected specimens from Thailand indicates that speci-mens with broad bands on the body and faint or nocheek dark marks are the xanthic form. HelmutDebelius provided information indicating hebelieved the two forms to represent a single speciesfrom observations in the field.Distribution. The species is widely distributed in

the Western Pacific (Western Australia and the GreatBarrier Reef in Australia, New Guinea, Brunei, Thai-land, Malaysia, the Philippines, Indonesia, Palau,China and Ryukyu Islands, Japan) and is known tobe associated with alpheid shrimps (Hayashi & Shi-ratori 2003). It occurs over a depth range of 1 to atleast 30 m.Relationships. Hoese & Larson (2004) recognised

five distinct species complexes of Cryptocentrus. Thegroups were based on various morphological andcolor characteristics. The Cryptocentrus cryptocentruscomplex is distinctive in having a bony ventral pro-jection from the operculum; the C. strigilliceps com-plex is distinctive in having transverse rows ofmandibular papillae and ctenoid scales on the body;the C. bulbiceps complex is distinctive in having awedge-shaped patch of predorsal scales and the headwith several oblique thin lines sloping backward andupward; the C. pavoninoides complex is distinctive inhaving 8-10 short rows of papillae radiating from theeye along the ventral and posteroventral margins ofthe eye onto the cheek; and the C. leucostictus com-plex is distinctive in having a very slender body, witha white stripe on the midline of the head, oftenextending onto the body. The various complexes pro-vide convenient grouping, but these may not neces-sarily represent monophyletic groups. The sixthgroup possessed no unifying characterists other thanlacking features of the other groups. The species inthis group include: Cryptocentrus caeruleopunctatus(Rüppell, 1830), Cryptocentrus cinctus (Herre, 1936),Cryptocentrus fasciatus (Playfair, 1866), Cryptocentrusgeniornatus Herre, 1935, Cryptocentrus leptocephalusBleeker, 1876, Cryptocentrus lutheri Klausewitz,1960, Cryptocentrus melanopus (Bleeker, 1860), Cryp-tocentrus shigensis Kuroda, 1956, Cryptocentrus sericusHerre, 1932 and Cryptocentrus yatsui Tomiyama,1936. All of the species in this group have similarmeristic features of 16-18 pectoral rays and longitu-dinal scale count of 60-80 and the head usually withsmall pearly white spots. The species described herefalls into this last group. Cryptocentrus sericus differsfrom all other species in the group, except C. fascia-tus, in having predorsal scales on the midline. Cryp-tocentrus sericus differs from C. fasciatus (see Fig. 6)

short horizontal thin black stripe above posteriorpreoperculum, sometimes absent; similar shorthorizontal stripe above operculum, contiguouswith lighter brown band crossing nape; cheek withnumerous scattered small light brown oval spots.Body with two vertical brown bands below firstdorsal fin, often with irregular margins and some-times partly interconnected and forming mottling;from second dorsal origin posteriorly 5 vertical ovalbrown spots present along midside, first justbehind first dorsal fin origin, second below middleof second dorsal fin, third below posterior end ofsecond dorsal fin, fourth anteriorly on caudalpeduncle and fifth just before base of caudal fin,spots elongating into bands extending from nearbase of second dorsal fin to base of anal fin in spec-imens above 55 mm SL; usually with very thinbrown vertical lines between broader body bands;fins dusky, pelvic fin with 3-5 dark bands infemales and 1-3 in males, often very faint in pre-served material, particularly in males.L i v e C o l o r a t i o n . Basic color pattern as

shown in Figures 1-3. Some underwater photos (Fig.5, Fishpix and photo from H. Debelius) show anoverall yellow or yellowish-orange coloration of thehead, body and fins. In these the dark spots on themidside are faint, but visible. Coloration of freshlycollected specimens differs little from the live col-oration. Yellow and banded color phases have beenobserved and collected on the Great Barrier Reef ofCryptocentrus cinctus Herre and these have beenobserved to change color (R. Kuiter, pers. comm.).Variation. Besides the live coloration variation and

sexual dimorphism mentioned above, the largestspecimens from Thailand and Hong Kong differslightly in having the first two dorsal spines moreelongate and some have the midside spots larger,appearing as bands in specimens above 55 mm SL. Inaddition the dark cheek spots and bars are faint orabsent in the largest specimens from Thailand and inone specimen from Hong Kong. These specimens,however, are the largest specimens examined and thedifferences are probably related to size. Photos of 54mm SL male from Thailand show a very similar col-oration to the specimen from the Great Barrier Reefshown here. Similarly, a 67 mm SL specimen fromWestern Australia has bands on the side narrowerthan in specimens from Thailand, but with the darkcheek bars and spots distinct. A 68 mm specimenfrom Sumatra has dark body bands as broad as inlarger specimens from Thailand, but with distinctdark cheek stripes and spots. Photos of freshly col-

Fig. 6. Cryptocentrus fasciatus from Great Barrier Reef. Photo by R. Kuiter.

Fig. 7. Cryptocentrus melanopus, male, ROM 73334, Vietnam. Photo by R. Winterbottom.

Fig. 8. Cryptocentrus melanopus, female, ROM 73334, Vietnam. Photo by R. Winterbottom.


169 aqua vol. 17 no. 3 - 10 July 2011

Fig. 10. Cryptocentrus leptocephalus, male, ROM 38763, Great Barrier Reef. Photo by R. Winterbottom.

Fig. 11. Underwater photo of Cryptocentrus leptocephalus, male, Koror, Palau. Photo by J. Sakaue.


170aqua vol. 17 no. 3 - 10 July 2011

sell, Fraser & Larson (2010) noted that Cryptocen-trus melanopus, described from a Castelnau paintingof a specimen from Singapore belongs to the groupand suggested that it was a senior synonym of Cryp-tocentrus leptocephalus. While further work is neededon the group, Cryptocentrus melanopus matches oneof the color forms with large red spots on the head;an elongate red stripe on upper pectoral fin base; abroad red bar from the ventral margin of the eye tothe middle of the upper jaw, rarely broken into twoparts; and the first dark body band below the seconddorsal fin sloping anteriorly ending entirely beforeanal fin, followed by 4 thin oblique bands. TheCastelnau painting shows only 3 oblique bands

in having three preopercular pores (vs. 2), narrowdark bars or oval spots along the midside (versus 5dark vertical bands extending from back to ventralsurface of body, often with irregular margins), bluecheek spots rounded (vs. obliquely elongated) andanterior margin of first dorsal fin elevated, with firsttwo dorsal spines the longest (vs. truncate torounded first dorsal margin with third or fourthspine the longest).

IDENTITY OF CRYPTOCENTRUS LEPTOCEPHALUS.It should be noted that there two color phases typ-

ically referred to as Cryptocentrus leptocephalus. Rus-

Fig. 12. Underwater photo of Cryptocentrus leptocephalus, female, Koror, Palau. Photo by J. Sakaue.


171 aqua vol. 17 no. 3 - 10 July 2011

a mixup in the specimens. However, it is more likelythat Herre’s drawing was actually a composite basedon the holotype and paratypes. Both species areincluded in the paratype series. Because of thisuncertainty and the confusion of separating the twospecies, we use the older name Cryptocentrus melano-pus here (Figs 7-9: note Fig. 9 is on the back cover).Cryptocentrus leptocephalus (including the holotypeRMNH 4665) has more numerous and smallerspots on the head; small round spots on upper pec-toral fin base; no broad bar below the eye to the jaws;and the first band below the second dorsal fin partlyoverlapping the anterior base of the anal fin, fol-lowed by three sloping bands (Figs 10-12). Males ofboth species have spots on the dorsal fins, whilefemales have red bands on the membranes betweenthe fin rays and spines.In Australia Cryptocentrus leptocephalus has been

collected from mangroves on coral reefs, while Cryp-tocentrus melanopus has been recorded from sandyreef flats, with algae. Johnson & Gill (2006)recorded and illustrated C. melanopus from the Gulfof Carpentaria (as C. leptocephalus). Both differ fromCryptocentrus sericus in having more anal rays (I,10),no predorsal scales and oblique dark bands on theside of the body.

behind the first band below the second dorsal fin,but agrees in other features with the species referredto here as Cryptocentrus melanopus. In some materialof Cryptocentrus melanopus from Australia some ofthe bands may fuse under the second dorsal fin,resulting in three bands on the body behind the sec-ond dorsal fin origin. Cryptocentrus melanopus hasalso been described as Cryptocentrus cheni Herre,1933 and Smilogobius obliquus Herre, 1934. Crypto-centrus melanopus has sometimes been identified asCryptocentrus singapurensis (Herre, 1936). The figureof the holotype given by Herre (1936) generallymatches C. melanopus in having the first band underthe second dorsal fin ending before the anal fin andin having a very dark area posteriorly with two orthree bands. However, the drawing shows smallspots as in C. leptocephalus. The specimen nowlabelled holotype (CAS SU 29807) is faded, butappears to have 4 bars beginning from the seconddorsal origin to the caudal peduncle, the body paleposteriorly and the first bar in contact with the analspine, as in Cryptocentrus leptocephalus. The currentnumber of apparent paratypes (CAS SU 16963, 35specimens; BMNH 1937.9.22, 1 specimen andZMH H.2864, 2 specimens) is greater than the 33listed by Herre and it is possible that there has been


172aqua vol. 17 no. 3 - 10 July 2011

Records of the Museums and Art Galleries of the Northern Ter-ritory 20: 167-174.

HOESE, D.F. & STEENE, R. (1978) Amblyeleotris randalli, anew species of gobiid fish living in association with alphaeidshrimps. Records of the Western Australian Museum 6 (4):379-389.

JOHNSON, J.W. & GILL, A. 2006. Reef and shore fishes ofSweers Island, Gulf of Carpentaria. Royal Geographical Soci-ety of Queensland, Geographical Monograph Series 10: 239-260.

KLAUSEWITZ, W. 1960. Fische aus dem Roten Meer. IV.Einige systematische und ökologisch bemerkenswerteMeergrundeln (Pisces, Gobiidae). Senckenbergiana Biologica41 (3/4): 149-162.

KUITER, R. H. & DEBELIUS, H. 2006. World Atlas of MarineFishes. IKAN-Unterwasserarchiv, Frankfurt, 720 pp.

KURODA, N. 1956. On an apparently new species of marinegoby of the genus Cryptocentrus. Annotationes ZoologicaeJaponensis. 29 (4): 242-245.

LARSON, H. K., JAAFAR, Z. & LIM, K. K. P. (2008). Anno-tated checklist of the gobioid fishes of Singapore. The Raf-fles Bulletin of Zoology 56 (12): 135-155.

LEVITON, A. E., GIBBS, R. H. HEAL, E. AND DAWSON, C. E.1985. Standards in Herpetology and Ichthyology: Part 1.Standard symbolic codes for institutional resource collec-tions in Herpetology and Ichthyology. Copeia 1985 (3):802-832.

MASUDA H., AMAOKA K., ARAGA C., UYENO T. & YOSHINO

T. (eds). 1984. The Fishes of the Japanese Archipelago. TokaiUniversity Press, Tokyo, 435 pp.

MYERS, R. F. 1999. Micronesian Reef Fishes. Coral Graphics,Guam, 330 pp.

OKAMURA, O. & AMAOKA, K. 1997. Sea Fishes of Japan.Yama Kei Publishers Co. Ltd., Tokyo, 783 pp.

PLAYFAIR, R. L. AND GÜNTHER, A. 1866. The Fishes of Zanz-ibar. London: John van Voorst.

RANDALL, J. E. 2004. Five new shrimp gobies of the genusAmblyeleotris from islands of Oceania. aqua, InternationalJournal of Ichthyology 8 (2): 61-78.

RANDALL, J. E., SHAO, K.-T. & CHEN, I.-P. 2003. A reviewof the Indo-Pacific gobiid fish genus Ctenogobiops, withdescriptions of two new species. Zoological Studies 42 (4):506-515.

RÜPPELL, W. P. E. 1830. Atlas zu der Reise im nördlichenAfrika. Fische des Rothen Meeres. 3 Frankfurt: H. L. Brün-ner: 95-141.

RUSSELL, B. C., FRASER, T. H. & LARSON, H. K. 2010.Castelnau’s collection of Singapore fishes described by PieterBleeker. The Raffles Bulletin of Zoology 58 (1): 93-102.

SHIBUKAWA, K. & SUZUKI, T. 2004. Vanderhorstia papilio, anew shrimp-associated goby from the Ryukyu Islands, Japan(Perciformes: Gobiidae, Gobiinae), with comments on thelimits of the genus. Ichthyological Research 51: 113-119.

SUZUKI, H., SUZUKI, T., SHIBUKAWA, Y. & YANO, K. 2004. APhotographic Guide to the Gobioid Fishes of Japan. Heibon-sha, Japan. 535 pp.

TOMIYAMA, I. 1938. Gobiidae of Japan. Japanese Journal ofZoology 7 (1): 37-112.

ACKNOWLEDGEMENTSBarry Russell kindly provided information and

photos of Castelnau’s painting of Cryptocentrusmelanopus. Bill Eschmeyer (CAS), Sue Morrison(WAM), Anthony Gill and James Maclaine(BMNH), Marinus Boeseman (RMNH), MadamBauchot (MNHN) and Katsusuke Meguro all pro-vided access to type and other material of Cryptocen-trus. Rick Winterbottom kindly provided three pho-tos used here and photos of freshly collected speci-mens of Cryptocentrus sericus from Thailand andPalau. Mark Mohlmann provided material of Cryp-tocentrus sericus from Thailand taken by Ukkrit Sat-apoomin. Rudie Kuiter and Gerald Allen collectedAustralian material. Roger Steene and Jack Randallprovided underwater photos of Cryptocentrus sericusfrom Australia, New Guinea and Palau and RogerSteene provided underwater photos of Cryptocentrusmelanopus. Helmut Debelius provided photos andinformation about Cryptocentrus sericus. HiroshiSenou kindly provided us with access to and use ofinformation from Fishpix.

REFERENCESBLEEKER, P. 1860. Derde bijdrage tot de kennis der vis-

chfauna van Singapoera. Natuurkundig Tijdschrift voor Ned-erlandsch-Indie 20: 446-356.

BLEEKER, P. 1876. Description de quelques espèces insulin-diennes inédites des genre Oxyurichthys, Paroxyurichthys etCryptocentrus. Verslagen en Mededeelongen der KoninklikjeAkademie van Wetenschappen, Letterkunde, en Schoone Kun-sten te Amsterdam (2) 9: 138-148.

HAYASHI, M., M. AIZAWA, M., ITOH, T. & ARAI, R. 1990,The marine gobiid fish fauna of Amami-Oshima Island,the Ryukyus, Japan. Memoires of the National ScienceMuseum (23): 123-150.

HAYASHI, M. & SHIRATORI, T. 2003. Gobies of JapaneseWaters. TBS Buritanica, Tokyo. 223 pp.

HERRE, A. W. 1932. Fishes from Kwangtung Province andHainan Island, China. Lingnan Science Journal 11: 423-443.

HERRE, A. W. 1933. Some Chinese gobies, with descriptionof one new species. Lingnan Science Journal 12: 429-430.

HERRE, A. W. 1934. Notes on Fishes in the Zoological Museumof Stanford University. 1. The fishes of the Herre PhilippineExpedition of 1931. The Newspaper Enterprise, HongKong, 106 pp.

HERRE, A. W. 1935. New fishes obtained by the CranePacific Expedition. Field Museum of Natural History, Zoo-logical Series Publication Number 335 18 (12): 383-438.

HERRE, A.W. 1936. Eleven new fishes from the Malay Penin-sula. Bulletin of the Raffles Museum, Singapore 12: 5-16.

HOESE, D. F. & LARSON, H. K. 2004. Description of a newspecies of Cryptocentrus (Teleostei: Gobiidae) from north-ern Australia, with comments on the genus. The Beagle,


Glossamia arguni, a new species of freshwater cardinalfish (Apogonidae) from West Papua Province, Indonesia

Renny K. Hadiaty1 and Gerald R. Allen2

1) Indonesian Institute of Sciences (LIPI), Research Centre for Biology, Jalan Raya Bogor, Km 46, Cibinong 16911, Indonesia. E-mail: [email protected]

2) Western Australian Museum, Locked Bag 49, Welshpool DC, Perth, Western Australia 6986. E-mail: [email protected]

Received: 11 March 2011 – Accepted: 12 June 2011

173 aqua vol. 17 no. 3 - 10 July 2011

Indonésie. Elle ressemble le plus à G. sandei qui occupeune large zone sud-centrale de la Nouvelle-Guinée, de laPurari River de Papouasie jusqu’au lac Yamur dans laPapua Province d’Indonésie. Les deux espèces se carac-térisent par des écailles relativement petites et un patron decoloration à barres. Pourtant, G. arguni a moins d’écaillesde la ligne latérale (43-45 contre 46-50) et cinq larges bar-res latérales contre 8-12 barres étroites pour G. sandei. Lanouvelle espèce compte aussi neuf rayons mous dorsauxcontre dix habituellement pour G. sandei.

SommarioUna nuova specie di pesce cardinale d’acqua dolce,

Glossamia arguni, è descritta sulla base di 13 esemplari di12.8-102.3 mm SL, raccolti nel novembre 2010 in torrentidel territorio dell’Arguni Bay nella regione del Bird’s Neck,Kaimana Regency, Papua Occidentale, Indonesia. Laspecie ad essa più simile è G. sandei, che è diffusa ampia-mente nella Nuova Guinea centromeridionale dal PurariRiver, Papua Nuova Guinea, al lago Yamur nella Provinciadi Papua, Indonesia. Entrambe le specie sono caratterizzateda scaglie relativamente piccole e da una livrea a barre. Tut-tavia, G. arguni n. sp. possiede un numero inferiore discaglie in linea laterale (43-45 versus 46-50) e cinqueampie barre sui lati invece di 8-12 sottili barre presenti inG. sandei. Inoltre, la nuova specie ha nove raggi molli dor-sali invece dei 10 riscontrati nella norma in G. sandei.

INTRODUCTIONThe cardinalfish family Apogonidae is best

known for its marine members, which includesmore than 300 species worldwide, mainly in trop-ical latitudes. It is one of the largest groups repre-sented in shallow coral reef habitats of the Indo-Pacific region. However, the genus Glossamia ofNew Guinea and northern Australia is entirelyrestricted to fresh waters. Allen (1991) provided akey to the species of Glossamia in addition to diag-

AbstractA new species of freshwater cardinalfish, Glossamia

arguni, is described on the basis of 13 specimens, 12.8-102.3 mm SL, collected in November 2010 from streamsin the Arguni Bay area of the Bird’s Neck region ofKaimana Regency, West Papua Province, Indonesia. It ismost similar to G. sandei, which ranges widely in south-central New Guinea from the Purari River of Papua NewGuinea to Lake Yamur in Papua Province of Indonesia.Both species are characterised by relatively small scales anda barred colour pattern. However, G. arguni n. sp. hasfewer lateral-line scales (43-45 vs. 46-50), and five broadbars on the sides compared with 8-12 narrow bars in G.sandei. The new species also has nine soft dorsal rays com-pared to the usual count of 10 for G. sandei.

ZusammenfassungBeschrieben wird eine neue Art eines Süßwasser-Kardinals:

Glossamia arguni, auf der Grundlage von 13 Exemplarenmit 12,8-102,3 mm SL, die im November 2010 in Bächender Gegend um die Arguni-Bucht, Region Bird’s Neck,Regentschaft Kaimana, Provinz West-Papua, Indonesien,gesammelt wurden. Die Exemplare dieser neuen Art ähnelnstark G. sandei, der im südlichen bis zentralen Neuguineaweit verbreitet ist, vom Purari-Fluss in Papua-Neuguinea biszum Yamur-See in der Provinz Papua in Indonesien. BeideArten sind durch relativ schmale Schuppen und Streifen-muster gekennzeichnet. Doch hat G. arguni n. sp. wenigerSeitenlinien-Schuppen (43-45 im Vergleich zu 46-50) undfünf breite Bänder auf den Flanken im Vergleich zu 8-12schmalen Streifen bei G. sandei. Außerdem zeigt die neueArt neun weiche Rückenflossenstrahlen, während es bei G.sandei normalerweise zehn sind.

RésuméUne nouvelle espèce de poisson cardinal d’eau douce,

Glossamia arguni, est décrite sur base de 13 spécimens, de12,8-102,3 mm de LS, collectés en novembre 2010 dansdes cours d’eau de la région de la baie d’Arguni du Bird’sNeck de la Kaimana Regency, West Papua Province,

Fig. 1. Satellite image of western New Guinea (Papua and West Papua provinces, Indonesia) and the Arguni Bay region (rec-tangular area outlined with white and shown in detail on left).

noses, illustrations, and distribution maps for sevenspecies recognized at that time. Allen et al. (2000)described an eighth species from the Timika regionof Irian Jaya (now Papua Province), Indonesia. The mountainous Central Range of New Guinea is

a formidable distributional barrier that prevents fau-nal interchange between northern and southerndrainages (Allen 1991). Northern species of Glos-samia include (approximate distributions in paren-theses): G. beauforti (Weber, 1907; Lake Sentani andpossibly Mamberamo River system), G. gjellerupi(Weber & de Beaufort, 1929; widespread northernNew Guinea), and G. wichmanni (Weber, 1907;coastal drainages of central north between PapuaNew Guinea border and Mamberamo River).Southern species include: G. aprion (Richardson,1842; widespread northern/eastern Australia andsouthern New Guinea from Fly River westward), G.narindica Roberts, 1978 (Bensbach and middle FlyRiver of south-western Papua New Guinea), G.sandei (Weber, 1907); Purari River, Papua NewGuinea to Etna Bay, Papua Province), G. timikaAllen et al., 2000 (Timika region of Papua Province,Indonesia), and G. trifasciata (Weber, 1913; FlyRiver, Papua New Guinea to Lorentz River, Papua

Province). Additionally, the Bird’s Head Peninsula,forming the western extremity of the island, has adistinctive fauna, at least at the specific level. How-ever, no members of Glossamia have been foundfrom the latter region despite extensive collectioneffort by the second author. The present paper describes a new species, the

ninth known member of the genus. It was collectedin November 2010 during an expedition led by theIndonesian Institute of Sciences (LIPI) to theArguni Bay region of the Bird’s Neck (Fig. 1), thenarrow isthmus of New Guinea situated betweenthe Bird’s Head Peninsula in the far west and themain body of the island. The Bird’s Neck area isamong the most rugged on the island of NewGuinea, containing extensive karst limestone,which forms a formidable barrier and makes inlandforays extremely difficult. Consequently, most ofthe region remains unexplored, with the exceptionof a few lakes and coastal streams. Previous expedi-tions by the first author resulted in a number ofnew discoveries, including endemic fishes from theTriton Lakes (Allen & Renyaan 1996a,b; Crowleyet al. 1995; Ivantsoff & Allen, 2011) and nearbyLake Mbutu (Allen 1998; Allen & Jenkins 1999).

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Fig. 2. Underwater photograph of Glossamia arguni,approximately 70 mm SL, Wainaga Stream, Arguni Bayregion, West Papua. Photo by G. R. Allen.

Fig. 3. Underwater photograph of Glossamia arguni,approximately 40 mm SL, Wainaga Stream, Arguni Bayregion, West Papua. Photo by G. R. Allen.

The present expedition, which was sponsored byConservation International in order to assist theKaimana government in an inventory of theircoastal and marine resources, focused predomi-nantly on the Arguni Bay region. Previously, Allen,Unmack & Hadiaty (2008) described Melanotae-nia ammeri (Melanotaeniidae) from a small creekflowing into Arguni Bay, near Gusimawa Village,but otherwise the fauna remained unknown untilour recent investigations. During the course of theweek-long expedition we collected fishes from twomain drainage areas of Arguni Bay including theKarora Estuary, an extensive Nipa palm embay-ment in the north and the Sewiki Basin in thesouth. The new Glossamia was found in streamhabitats throughout this area.

MATERIALS AND METHODSProportional measurements of type specimens

are presented in Table I as percentages of the stan-dard length. Counts and measurements that appearin parentheses refer to the range for paratypes ifdifferent from the holotype. Type specimens aredeposited at Museum Zoologicum Bogoriense,Cibinong, Indonesia (MZB) and the Western Aus-tralian Museum, Perth (WAM).Lengths given for specimens are standard length

(SL), measured from the front of the upper lip tothe base of the caudal fin (posterior end of hypuralplate); body depth is the maximum depth from thebase of the dorsal spines; body width is measuredjust posterior to the gill opening; head length istaken from the front of the upper lip to the end ofthe opercular membrane, and snout length fromthe same anterior point to the fleshy edge of theorbit; orbit diameter is the greatest fleshy diameter,and interorbital width the least bony width; caudalpeduncle depth is the least depth, and caudalpeduncle length the horizontal distance betweenverticals at the posterior base of the anal fin and thebase of the caudal fin; lengths of fin spines and softrays are measured to their extreme bases; caudalconcavity is the horizontal distance between verti-cals at the tips of the shortest and longest caudal finrays; pectoral ray counts include the upper rudi-mentary ray; lateral line scale counts are made tothe base of the caudal fin (hence do not include thepored scales posterior to the hypural plate); gillraker counts are made on the first gill arch andinclude developed rakers only (those which arehigher than the width of their base); the count ofthe upper-limb raker is given first, followed by the

lower limb count; the raker at the angle is con-tained in the lower limb count.

Glossamia arguni n. sp.(Figs 2-5; Table I)

Holotype: MZB 19771, male, 89.4 mm SL,Wahisewar River, 03°03’55.1”S, 133°58’18.8”E,tributary of Tongaran River, about 11 km south-east of Gusimawa Village, Arguni Bay region, WestPapua Province, Indonesia, seine, R. Hadiaty andparty, 2 November 2010.Paratypes (12 specimens, 12.8.0-102.3 mm SL):

MZB 19611, 3 specimens, 18.5-64.0 mm SL, col-lected with holotype; MZB 19648, 5 specimens,12.8-47.7 mm SL, small tributary, 03°09’26.0”S;133°57’05.4”E, of Waronais River, near WainagaVillage, about 16 km south-east of Gusimawa Vil-lage, Arguni Bay region, West Papua, Indonesia,seine, R. Hadiaty and party. 3-4 November 2010;MZB 19692, 2 specimens, 24.1-26.1 mm SL,Buguma stream, 03°15’03.9”S; 133°47’40.4”E,tributary of Sewiki River, 6.6 km north of Lake

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Renny K. Hadiaty and Gerald R. Allen

Fig. 4. Glossamia arguni, preserved male, holotype, MZB19771, 89.4 mm SL, Arguni Bay region, West Papua,Indonesia. Photo by G. R. Allen.

Fig. 5. Glossamia arguni, preserved paratype, 64.0 mm SL,Arguni Bay region, West Papua, Indonesia. Photo by G. R.Allen.

Table I. Proportional measurements of selected type specimens of Glossamia arguni expressed as percentage of the standardlength.

Holotype Paratype Paratype Paratype Paratype ParatypeMZB WAM P. WAM P. MZB MZB MZB19771 33339-001 33339-001 19611 19648 19648

Standard length (mm) 89.4 102.3 72.0 64.0 41.6 40.9Body depth 35.5 36.5 33.9 35.9 36.1 35.7Body width 17.6 18.9 15.8 18.6 15.4 14.2Head length 39.0 37.5 38.3 37.8 38.5 38.1Snout length 9.4 11.8 10.7 10.9 10.1 9.3Eye diameter 10.2 10.0 10.7 11.7 13.2 12.5Bony interorbital width 8.3 7.9 7.9 8.0 9.1 7.8Upper jaw length 21.4 20.0 20.6 22.3 21.2 21.5Depth of caudal peduncle 15.0 15.2 14.3 16.3 15.1 13.7Length of caudal peduncle 24.5 24.0 24.6 25.3 28.4 27.1Predorsal distance 49.3 45.5 47.8 44.1 50.0 47.2Preanal distance 65.8 65.5 62.1 64.5 62.7 62.6Prepelvic distance 39.6 43.5 38.3 37.7 38.5 43.01st dorsal spine length 4.6 5.5 5.0 5.8 7.5 7.82nd dorsal spine length 18.6 17.9 21.0 20.2 20.7 20.53rd dorsal spine length 3.8 4.4 4.4 3.3 2.9 5.11st spine of 2nd dorsal fin length 11.6 11.4 11.7 11.9 11.5 13.2Longest soft dorsal ray 17.2 16.6 19.2 17.7 17.1 17.61st anal spine length 3.7 4.5 4.2 3.8 5.0 4.92nd anal spine length 12.5 12.3 12.6 13.1 15.4 15.2Longest soft anal ray 16.6 16.8 18.9 16.7 20.4 21.5Caudal fin length 28.0 26.5 28.9 27.3 30.5 30.1Caudal concavity 4.0 4.8 5.0 6.4 7.5 6.1Pectoral fin length 22.6 20.6 22.6 20.2 18.5 19.6Pelvic fin spine length 13.6 12.9 14.0 13.1 13.5 15.4Pelvic fin length 20.2 19.6 21.1 18.3 22.8 24.0

Sewiki, Arguni Bay region, West Papua, Indonesia,seine, R. Hadiaty and party, 5 November 2010;WAM P.33339-001, 2 specimens, 72.0-102.3 mmSL, collected with holotype. Diagnosis: Dorsal rays VI-I,9; anal rays II,8; pec-

toral rays 15; developed gill rakers on first arch 1 +6; lateral-line scales 43-45 (usually 44); greatestbody depth 2.7-3.0 in SL; live colour generally palegrey grading to greyish brown dorsally with five

broad dark brown bars on side from level of pectoralfin to anterior part of caudal peduncle; bars more orless merging dorsally with broad, irregular stripe onupper side extending from upper rear margin ofoperculum to middle of caudal-fin base; saddle-likemarking (abbreviated bar) ventrally on posteriorcaudal peduncle; smaller fish (under about 40 mmSL) with more prominent brown bars on side,noticeably wider than pale grey interspaces.

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Fig. 6. Aquarium photograph of Glossamia sandei, approximately 100 mm SL, Timika, Papua Province, Indonesia. Photo byG. R. Allen.

Fig. 7. Aquarium photograph of Glossamia timika, approximately 90 mm SL, Timika, Papua Province, Indonesia. Photo byG. R. Allen.

Description: Values in parentheses refer toparatypes (Table I). Dorsal rays VI-I,9; anal raysII,8; all dorsal and anal soft rays branched, last tobase; pectoral rays 15, 1-2 lowermost and twouppermost unbranched; pelvic rays I,5, all raysbranched; principal caudal rays 17, uppermost andlowermost rays unbranched; lateral-line scales 44(43-45, usually 44), plus 2-3 pored scales posteriorto hypural plate; scales above lateral line to originof dorsal fin 4; scales below lateral line to origin ofanal fin 12; median predorsal scales 12; circumpe-

duncular scales 21 (21-22); developed gill rakers 1+ 6, usually 2-3 very low, barely detectable rudi-ments at beginning of upper limb and 3-4 similarrudiments at end of lower limb of adults; bran-chiostegal rays 7.Body depth 2.8 (2.7-3.0) in SL; body width 2.0

(1.9-2.5) in depth; head length 2.6 (2.6-2.7) in SL;dorsal profile of head concave (concavity more pro-nounced in large specimens), snout rounded; snoutlength 4.2 (3.2-4.1) in head; orbit diameter 3.8(2.9-3.8) in head; interorbital width 4.7 (4.2-4.9)

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in head; caudal peduncle depth 2.6 (2.3-2.8) inhead; caudal peduncle length 1.6 (1.4-1.6) in head.Mouth large, maxilla reaching a vertical through

rear edge of eye or slightly posterior of this point,upper jaw length 1.8 (1.7-1.9) in head; mouthoblique, gape forming angle of about 30 degrees tohorizontal axis of head and body; posterior edge ofmaxilla straight; no supramaxilla present; lower jawslightly inferior; upper jaw with band of small vil-liform teeth in about 10-12 irregular rows withtoothless gap at front of jaw; lower jaw with bandof similar teeth in about 6-8 irregular rows; irregu-lar row of small teeth forming V-shape on vomerand straight, narrow row on each palatine; tonguebroad based with rounded tip.Anterior nostril small, low-rimmed, membranous

tube, about level with lower edge of pupil and posi-tioned midway between anterior edge of eye andupper lip; posterior nostril ovate without rim, online connecting anterior nostril to top of pupil,more than twice as large as anterior nostril;cephalic lateralis pores numerous, especially oninterorbital, around eye, and on ventral surface oflower jaw, mainly arranged in transverse and longi-tudinal series; preopercular margin and preopercu-lar ridge smooth.Scales weakly ctenoid; lateral line conspicuous,

nearly paralleling dorsal contour of body and end-ing at caudal-fin base (2-3 pored scales posterior tohypural); no scales on dorsal and anal fins except

low sheath at base of second dorsal and anal fins;small scales at base of caudal fin; no scales at baseof pectoral fins; small scales covering basal areabetween pelvic fins.Origin of first dorsal fin above eighth or ninth lat-

eral-line scale; first dorsal spine slender and short,8.5 (4.0-7.7) in head; second and third dorsal spinelongest, 2.1 (1.8-1.9) in head; middle soft dorsalrays longest, 2.3 (2.0-2.3) in head; origin of analfin even with origin of second dorsal fin; first analspine very short, 8.5 (7.6-10.1) in head; secondanal spine 3.1 (2.5-3.0) in head; anterior anal softrays longest, 2.4 (1.8-2.3) in head; caudal fin emar-ginate with rounded lobes, its length 1.4 (1.3-1.4)in head; caudal concavity 9.7 (5.2-7.7) in head;upper pectoral rays longest, 1.7 (1.7-2.1) in head;origin of pelvic fins even with pectoral fin base;first pelvic soft ray longest, reaching to about anusor slightly beyond, its length 1.9 (1.6-2.1) in head.C o l o u r i n l i f e (from underwater pho-

tographs, Fig. 2): generally pale grey grading togreyish brown dorsally with five broad dark brownbars on side from level of pectoral fin to anteriorpart of caudal peduncle; bars more or less mergingdorsally with broad, irregular stripe on upper sideextending from upper rear margin of operculum tomiddle of caudal-fin base; saddle-like marking(abbreviated bar) ventrally on posterior caudalpeduncle; small (about one-half to one-third pupilsize) blackish spot on upper preopercle, just behindeye; short yellowish stripe across middle of opercu-lum with dark brownish area just below; finstranslucent greyish to blue grey without distin-guishing marks. Smaller fish (under about 40 mmSL, Fig. 3) with more prominent brown bars onside, noticeably wider than pale grey interspaces. C o l o u r o f h o l o t y p e i n a l c o h o l (Fig.

4): head and body generally brownish with poorlydefined series of five darker brown bars on sidefrom level of pectoral fin to anterior caudal pedun-cle; lips tannish; fins dusky brown to tan.C o l o u r o f p a r a t y p e s i n a l c o h o l : inter-

mediate-sized fish (64 mm SL, Fig. 5) overall paletan to light brown with series of six, bold darkbrown bars, including two narrower bars on caudalpeduncle; first bar below first dorsal fin origin, sec-ond below rear part of first dorsal fin, third belowanterior part of second dorsal fin, and fourth belowrear part of second dorsal fin; oblique brown bandangling dorsally from upper rear corner of eye andhint of brown band from lower rear corner of eye tojust behind maxillary; fins dusky brownish to tan-

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Fig. 8. Glossamia arguni, yolk-sac embryo, 4 mm TL, approx-imately 3-6 hours after hatching. Photo by R. Hadiaty.

nish; middle of pelvic fin with dusky brown spot. In some paratypes the two bars on the caudal

peduncle are linked dorsally and ventrally with anarrow stripe, forming a rectangular marking. Thelargest specimen (102.3 mm SL) faded immedi-ately after death and is uniformly greyish brown inpreservative, without markings.Remarks: Glossamia arguni is distinguished from

other members of the genus by the combination offeatures indicated in the key to species that appearsbelow. It is most similar to G. sandei (Weber, 1907)(Fig. 6), which ranges widely in south-central NewGuinea from the Purari River of Papua NewGuinea to Lake Yamur in Papua Province ofIndonesia. Both species are characterised by rela-tively small scales and a barred colour pattern.However, G. arguni has fewer lateral-line scales(43-45 versus 46-50) and five broad bars on thesides compared with 8-12 narrow bars in G. sandei.The new species also has nine soft dorsal rays com-pared to the usual count of 10 for G. sandei. Glos-samia timika (Allen, Hortle, and Renyaan, 2000)from south-western Papua Province, Indonesia,also possesses a barred pattern (Fig. 7) consisting ofthree primary dark bars on the side with three nar-row bars between the first two main bars. It alsodiffers in having only 24-26 lateral-line scales.

Key to the species of Glossamia from southernNew Guinea

1a. Body with 5-13 dark vertical bars (sometimesfaint in large adults) ..................................... 2

1b. Body with 3-4 irregular, dark vertical bars, ormottled brown ............................................. 4

2a. Lateral-line scales 24-26; 8 gill rakers on firstarch; anal rays II,9; 7 dark bars on side...................................................... Glossamia timika

2b. Lateral-line scales 43-50; 7 gill rakers on firstarch; anal rays II,8; 5-12 dark bars on side .. 3

3a. Body with 5-6 dark bars; lateral line scales 42-45; soft dorsal rays 9........... Glossamia arguni

3b. Body with 8-12 dark bars; lateral-line scales46-50; soft dorsal rays 10............................................................................ Glossamia sandei

4a. First gill arch usually with 8 gill rakers; darkbars distinct, rarely obscured by uniform darkcolour, but never by mottling; dark bar belowfirst dorsal fin wider than eye................................................................ Glossamia trifasciata

4b. First gill arch with 7 gill rakers, dark bars some-times obscured by mottling, but never by uni-

form dark colour; dark bar below first dorsalfin much narrower than eye ......................... 5

5a. Body usually with strong mottling betweendark bars; snout immediately below posteriornostril without distinctive mark; body depth atfirst dorsal origin 2.7-3.0 in SL.................................................................. Glossamia aprion

5b. Body without mottling between dark bars;snout immediately below posterior nostril withdistinct dark spot; body depth at first dorsalorigin 3.2-3.5 in SL ....... Glossamia narindica

Behaviour and Ecology: The new species wasobserved underwater with the use of scuba gear bythe second author in a small (about 3-5 m wide andpools to 2.5 m depth), clear stream flowing throughlowland rainforest habitat. Widely scattered, solitaryindividuals were seen hovering in sheltered loca-tions, frequently among limestone boulders or theroots and branches of fallen trees. The stream con-tained pure fresh water (temperature 24-25 C), butwater levels were influenced by tidal fluctuation.Co-inhabitants of the stream included (in order ofabundance) Gobiopterus species, Redigobius chryso-soma (Bleeker), Melanotaenia ammeri Allen,Unmack & Hadiaty, Hypseleotris species, Scatopha-gus argus (Linnaeus), Oxyeleotris aruensis (Weber),Toxotes chatareus (Hamilton) and Leptachirus alleniRandall. Additional co-inhabitants at other loca-tions where G. arguni was collected included Pseudo-mugil species, Microphis brevidorsalis (de Beaufort)and the following gobioid fishes: Glossogobiusspecies, Gobiopterus species, Periophthalmus weberiEggert, Mugilogobius mertoni (Weber), M. rivulusLarson, Giuris margaritacea (Valenciennes) Mo gurn -da species, Oxyeleotris aruensis (Weber), O. fimbriata(Weber) and O. nullipora Roberts.One of the large adult male specimens of G. arguni

that was collected had a batch of 63 eggs in itsmouth. These were placed in a small plastic bag witha few aquatic plants. Six hours later it was noticedthat some of the eggs had hatched and a photographof the yolk-sac embryo is presented as Fig. 8. Theegg-brooding habits of marine apogonids was sum-marized by Allen (1975 and 1993). However, thereis a general lack of knowledge concerning the fresh-water Glossamia species. Allen et al. (2008) reportedthat in at least one species, G. trifasciata, the malecontinues to shelter its young in the mouth for anundetermind period after hatching.Distribution: Glossamia arguni is known only

from freshwater streams in the Arguni Bay region

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of West Papua Province, Indonesia. This locationrepresents the farthest west record for the genus onthe island of New Guinea and is situated approxi-mately 130 km north-west of Lake Yamur, the pre-vious limit of distribution. No species of Glossamiawere collected during surveys by the second authorat intervening locations including the Etna Bayand Triton Lakes regions.Etymology: The new species is named arguni

with reference to the type locality, which is situatedin the Arguni Bay region.

ACKNOWLEDGEMENTSWe are greatly indebted to Richard Schneider and

Fabian Oberfeld from Los Angeles, California,who provided financial sponsorship, excellentcompanionship, and field assistance. Special thanksare reserved for the local government and themasyarakat adat (traditional communities) ofKaimana and Arguni Bay for allowing us to visittheir spectacularly beautiful homeland and fortheir commitment to preserve it. We also thank ourLIPI colleagues Daisy Wowor, Ristiyanti Marwotoand Mulyadi, and Samuel Renyaan (University ofCendrawasih) for their advice and collecting assis-tance. Max Ammer provided valuable collectingassistance. Ken and Josephine Wiedenhoeft andtheir crew on Puti Raja were excellent hosts andprovided essential logistic assistance. We are alsoindebted to Nimrod (Roy) Tafre from the KaimanaSpatial Planning Department and Zeth Parindingfrom the Kaimana Department of Nature Conser-vation for their help in making our expedition pos-sible and collection assistance. The staff of theKaimana office of Conservation International (CI)were very helpful with planning issues and com-munication with local authorities. Finally, we areespecially grateful to Mark V. Erdmann (CI) for histireless organizational efforts and essential assis-tance during the Arguni Bay expedition.

REFERENCESALLEN, G. R. 1975. The biology and taxonomy of the car-

dinalfish, Sphaeramia orbicularis (Pisces: Apogonidae).Journal of the Royal Society of Western Australia 58 (3): 86-92.

ALLEN, G. R. 1991. Field Guide to the Freshwater Fishes ofNew Guinea. Christensen Research Institute, Madang,Papua New Guinea. 268 pp.

ALLEN, G. R. 1993. Cardinalfishes (Apogonidae) ofMadang Province, Papua New Guinea, with descriptionsof three new species. Revue française d’Aquariologie Herpé-tologie 2 (1): 9-20.

ALLEN, G. R. 1998. A new genus and species of rainbowfish(Melanotaeniidae) from fresh waters of Irian Jaya, Indone-sia. Revue Française Aquariologie 2 (1-2): 11-16.

ALLEN, G. R., HORTLE, K. & RENYAAN, S. J. 2000. Fresh-water Fishes of the Timika Region, New Guinea. P.T.Freeport Indonesia, Timika, Indonesia. 175 pp.

ALLEN, G. R. & JENKINS, A. P. 1999. Two new species ofMogurnda (Osteichthyes: Eleotrididae) from the Etna Bayregion, Irian Jaya, Indonesia. Ichthyological Exploration ofFreshwaters 10 (3): 237-246.

ALLEN, G. R. & RENYAAN, S. J. 1996a. Three new species ofrainbowfishes (Melanotaeniidae) from the Triton Lakes,Irian Jaya, New Guinea. Aqua, Journal of Ichthyology andAquatic Biology 2 (2): 13-24.

ALLEN, G. R. & RENYAAN, S. J. 1996b. Eleotrid fishes of theTriton Lakes, Irian Jaya, with descriptions of four newspecies. Revue Française Aquariologie 23 (1-2): 47-56.

ALLEN, G. R., STOREY, A. W. & YARRAO, M. 2008. Fresh-water Fishes of the Fly River Papua New Guinea. Ok TediMining, Tabubil, Papua New Guinea. 216 pp.

ALLEN, G. R., UNMACK, P. J. & HADIATY, R. K. 2008. Twonew species of rainbowfishes (Melanotaeniidae: Melanotae-nia) from western New Guinea (Papua Barat Province,Indonesia). aqua, International Journal of Ichthyology 14(4): 209-224.

CROWLEY, L. E. L. M., IVANTSOFF, W. & ALLEN, G. R.1995. Description of a new species of hardyhead, Cratero-cephalus fistularis (Pisces: Atherinidae) from Irian Jaya.Records of the Western Australian Museum 17 (3): 325-329.

IVANTSOFF, W. & ALLEN, G. R. 2011. A new species andgenus of a large and unusual freshwater hardyhead,Sashatherina giganteus (Pisces: Atherinidae) from WestPapua, Indonesia and a comparison with its closest rela-tives of the genus Craterocephalus. aqua International Jour-nal of Ichthyology 17 (1): 43-57.

RICHARDSON, J. 1842. Contributions to the ichthyology ofAustralia. Annals and Magazine of Natural History (NewSeries) 9 (55): 15-31.

ROBERTS, T. R. 1978. An ichthyological survey of the FlyRiver in Papua New Guinea with descriptions of newspecies. Smithsonian Contributions to Zoology 281: 1-72.

WEBER, M. 1907. Süsswasserfische von Neu-Guinea einBeitrag zur Frage nach dem früheren Zusammenhang vonNeu-Guinea und Australien. In: Nova Guinea. Résultats del’expédition scientifique Néerlandaise à la Nouvelle-Guinée.Süsswasserfische von Neu-Guinea ein Beitrag zur Frage nachdem früheren Zusammenhang von Neu-Guinea und Aus-tralien 5 (Zoology.), part 2: 201-267.

WEBER, M. 1913. Süsswasserfische aus Niederländisch Süd-und Nord-Neu-Guinea. In: Nova Guinea. Résultats de l’ex-pédition scientifique Néerlandaise à la Nouvelle-Guinée.Zoologie. Leiden. Süsswasserfische aus Niederländisch Süd-und Nord-Neu-Guinea 9 (4): 513-613, Pls. 12-14.

Weber, M. & de Beaufort, L. F. 1929. The fishes of the Indo-Australian Archipelago. Volume 5. Anacanthini, Allotriog-nathi, Heterostomata, Berycomorphi, Percomorphi: familiesKuhliidae, Apogonidae, Plesiopidae, Pseudoplesiopidae, Pria-canthidae, Centropomidae. E. J. Brill Ltd., Leiden. 458 pp.

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BLABER, S. J. M. 1980. Fish of the Trinity inlet systemof North Queensland, with notes on the ecology offish faunas of tropical Indo-Pacific estuaries. Aus-tralian Journal of Marine and Freshwater Research31:137-46.

DAY, J. H., BLABER, S. J. M., & WALLACE, J. H. 1981.Estuarine fishes. In: Estuarine Ecology with Particu-lar Reference to Southern Africa. (Ed. J.H. Day.):197-221. A. A. Balkema, Rotterdam.

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TREWAVAS, E. 1983. Tilapiine Fishes of the GeneraSarotherodon, Oreochromis and Danakilia. BritishMuseum (Natural History), London, 583 pp.

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aquaInternational Journal of Ichthyology

Vol. 17 (3), 10 July 2011

Contents:

Leonardo A. Abitia-Cárdenas, Xchel G. Moreno-Sánchez*, Deivis S. Palacios-Salgado and Ofelia Escobar-Sánchez: Feeding habits of the convict surgeonfish Acanthurus triostegus(Teleostei: Acanthuridae) on the Los Frailes reef, Baja California Sur, Mexico .......................................121-126

Richard Winterbottom: Six new species of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature ....................127-162

Doug Hoese, Koichi Shibukawa and Jiro Sakaue: A redescription of the gobiid fish Cryptocentrus sericus Herre, with clarification of Cryptocentrus leptocephalus and C. melanopus ...............163-172

Renny K. Hadiaty and Gerald R. Allen: Glossamia arguni, a new species of freshwater cardinalfish (Apogonidae) from West Papua Province, Indonesia .............................................................................173-180

Papers appearing in this journal are indexed in: Zoological Record; BioLIS – Biologische Literatur Information Senckenberg;

www.aqua-aquapress.com; www.aquapress-bleher.com; www.Joachim-Frische.com

Cover photo: Left lateral view of Trimma erdmanni (live), Kawe Island, Raja Ampat. Photo by M. V. Erdmann.

Fig. 9. Underwater photo of Cryptocentrus melanopus, female, Bali, Indonesia. Photo by R. Steene. See pages 155-162.

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