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1 aqua, International Journal of Ichthyology Six new species of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat 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. rickw@rom.on.ca Received: 30 December 2010 Accepted: 01 April 2011 Abstract Recent (2010) fieldwork in the Raja Ampat Islands of Indonesia resulted in the collection of seven undescribed species of Trimma, six of which are described here. Two of these are currently only known from these islands, the others are known from other parts of the western Pacific. Trimma cheni n. sp. is distinguished from other species of Trimma in having 8-9 scales in the predorsal midline, 2-3 scales on the opercle, an elongate second dorsal spine reaching posteriorly to the bases of rays 1-4 of the second dorsal fin, middle pectoral fin rays branched, fifth pelvic fin ray branched once, a dark basal stripe in the dorsal fins, scale pockets indistinctly outlined with darker pigment, and, in life, two red to orange bars across the cheek and three diffuse yellow stripes on the body (most obvious along the caudal peduncle). It has been recorded from Palau, the Philippines, and Sulawesi and Flores in Indonesia. Trimma erdmanni n. sp. has a reddishorange body with a dark red lateral stripe with darker borders on the body which extends anteriorly onto the head, where it bifurcates, and there is a thin longitudinal light stripe below the eye. It usually lacks scales in the predorsal midline, the second spine of the first dorsal is elongated, there are 9 dorsal and 8 anal rays, a single branch in the fifth pelvic fin ray, and gill rakers on the first gill arch. The species is known from the Raja Ampat islands, the Hermit Islands and Madang (Papua New Guinea), and from photographs from El Nido, Palawan Island and Davao Gulf, Mindanoro, Philippines. Trimma habrum n. sp. has a bony interorbital width equal to the pupil diameter, 8-9 scales in the predorsal midline, 14 unbranched pectoral fin rays, an unbranched fifth pelvic fin ray, no basal membrane joining the fifth pelvic fin rays across the midline, usually a single full row of cheek scales, and scales on the upper two-thirds of the opercle. The fresh colouration is diagnostic: a pale translucent dorsum with light yellow blotches and the base of each element of the dorsal fin surrounded by a red spot, a thin red bar along the posterior margins of the hypurals, no dark pigment at all on the hypural region of the peduncle, and a darkly pigmented covering to the dorsal margins of the swim bladder, neural sheath, and the brain. It is currently known only from a single collection made at Kerou Island, Fam Islands in Raja Ampat, Indonesia. Trimma haimassum n. sp. is characterized by a relatively deep body, numerous irregular scales with about 30 lateral rows and over 15 scales in the anterior transverse series, a fifth pelvic fin ray that branches twice dichotomously, usually a somewhat elongate second dorsal spine, no predorsal, cheek or opercular scales, a moderately wide bony interorbital with a fleshy median ridge between the eyes, and a dermal ridge anterior to the first dorsal spine. There is a red spot or elongate blotch above the opercle in live and fresh material, and the dorsal surface of the snout has a reticulated dark pattern, with dark transverse stripes over the dorsal margin of the orbit. The species is known from south-western Sulawesi north to Palawan and eastwards to the Solomon Islands. Trimma papayum n. sp. is unique among the described species of the genus in having a one-third pupil diameter black ocellated spot on and just behind the fourth dorsal fin spine. It has 9 dorsal and 8 anal fin rays, a single branch point in the fifth pelvic fin ray, anterior and 8-9 posterior transverse scale rows, 5-10 scales in the predorsal midline, and a single row of 3 scales on the upper margin of the opercle. Freshly collected specimens are orange red in overall colouration, with scattered diffuse yellow spots. The species is known only from Indonesia, at Maumere, Flores and Kawe Island, Raja Ampat. Trimma xanthochrum n. sp. is characterized by a wide interorbital region (80-100% pupil diameter), a second dorsal spine usually reaching posteriorly to between the bases of the second to third dorsal fin rays, pectoral rays usually with 7-8 branched rays, vertical rows of sensory papillae below eye of 2-3 papillae in rows 1-4 and 4-5 in row 5, a caudal blotch which has a lower half about two-thirds the width of the upper half, and usually an overall yellowish body with yellow at least proximally in the caudal fin. It is currently known with certainty only from the Raja Ampat islands. Zusammenfassung Bei einer neueren Forschungsreise zum Raja-Ampat-Archipel Indonesiens (2010) wurden sieben unbeschriebene Trimma-Arten entdeckt, von denen sechs hier beschrieben werden. Zwei der neuen Arten kommen nach heutiger Kenntnis nur an diesen Inseln vor, die anderen sind auch von anderen Teilen des Westpazifiks bekannt. Trimma cheni n. sp. unterscheidet sich von den anderen Trimma-Arten durch fol- 127 aqua vol. 17 no July 2011

2 Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature gende Merkmale: 8-9 Schuppen auf der prädorsalen Mittellinie, 2-3 Schuppen auf dem Kiemendeckel, einen verlängerten zweiten Rückenstachel, der nach hinten bis zur Basis der Flossenstrahlen 1-4 der zweiten Rückenflosse reicht, verzweigte mittlere Strahlen der Brustflosse, einmal verzweigten fünften Bauchflossenstrahl, einen dunklen Streifen am Grund der Rückenflossen, undeutlich mit dunklerem Farbstoff umrahmte Schuppentaschen, außerdem beim lebenden Tier zwei rote bis orangefarbene Bänder über die Wange hinweg und drei schwach gelbe Streifen auf dem Rumpf (am deutlichsten sichtbar auf dem Schwanzstiel). Diese Art konnte bei Palau, den Philippinen und Sulawesi und an der Insel Flores in Indonesien nachgewiesen werden. Trimma erdmanni n. sp. hat einen rötlich orangefarbenen Rumpf und einen dunkelroten Seitenstreifen mit dunkleren Rändern auf dem Rumpf, der sich nach vorne bis zum Kopf erstreckt, wo er sich gabelt, außerdem einen dünnen hellen Längsstreifen unter dem Auge. Normalerweise fehlen hier Schuppen auf der prädorsalen Mittellinie, der zweite Stachel der ersten Rückenflosse ist verlängert, die Rückenflossen haben 9 Flossenstrahlen, die Afterflosse 8, der fünfte Bauchflossenstrahl verzweigt sich einmal, und der erste Kiemenbogen trägt Kiemenreusen. Fundorte dieser Art sind die Inseln Raja Ampat, die Hermit-Inseln und Madang (Papua-Neuguinea) sowie, durch Fotos belegt, El Nido, Palawan-Insel und der Davao-Golf, Mindanoro, Philippinen. Bei Trimma habrum n. sp. entspricht die Breite des Interorbitalknochens dem Pupillendurchmesser, Vertreter dieser Art haben 8-9 Schuppen auf der prädorsalen Mittellinie, 14 unverzweigte Brustflossenstrahlen, einen unverzweigten fünften Bauchflossenstrahl, keine Basalmembran am fünften Bauchflossenstrahl über die Mittellinie hinweg, normalerweise eine einzelne volle Reihe von Wangenschuppen und Schuppen auf dem oberen Zweidrittel des Kiemendeckels. Außerdem ist die Färbung lebender oder frischtoter Exemplare artkennzeichnend: der Rücken blass durchscheinend mit hellgelben Flecken, die Basis von jedem Element der Rückenflosse von einem roten Fleck umrahmt, ein dünner roter Streifen entlang den hinteren Rändern der Hypurale, überhaupt kein dunkles Pigment in der hypuralen Gegend des Schwanzstiels sowie eine dunkel pigmentierte Bedeckung der dorsalen Ränder der Schwimmblase, der Nervenscheide und des Gehirns. Bisher ist diese neue Art nur von einem einzigen Forschungsfang an der Kerou-Insel, Fam-Inseln im Raja-Ampat-Archipel, Indonesien, bekannt. Trimma haimassum n. sp. ist gekennzeichnet durch einen relativ tiefen Rumpf, zahlreiche unregelmäßige Schuppen mit etwa 30 seitlichen Reihen und über 15 Schuppen in der vorderen Querreihe, einen fünften Bauchflossenstrahl, der sich zweimal dichotom verzweigt, normalerweise einen etwas verlängerten zweiten Rückenstachel, das Fehlen von Schuppen prädorsal, an der Wange und auf dem Kiemendeckel, einen mäßig breiten Interorbitalknochen mit einem fleischigen medianen Grat zwischen den Augen sowie eine Hautleiste vor dem ersten Rückenstachel. Bei lebenden oder frischtoten Exemplaren zeigt sich in roter Farbe ein Punkt oder länglicher Fleck oberhalb vom Kiemendeckel, und die rückwärtige Oberfläche der Schnauze weist ein netzartiges dunkles Muster auf, mit dunklen Querstreifen über dem rückwärtigen Rand der Augenhöhle. Man fand diese Art am südwestlichen Sulawesi nördlich von Palawan und nach Osten zu an den Salomoninseln. Trimma papayum n. sp. zeichnet sich unter den beschriebenen Arten dadurch aus, dass sich auf und direkt hinter dem vierten Rückenflossenstachel ein schwarzer Augenfleck mit dem Durchmesser eines Drittels der Pupille befindet. Außerdem haben Vertreter dieser Art 9 Rückenflossen- und 8 Afterflossenstrahlen, nur einen Verzweigungspunkt beim fünften Bauchflossenstrahl, vordere und 8-9 hintere quer laufende Schuppenreihen, 5-10 Schuppen auf der prädorsalen Mittellinie und eine einzelne Reihe von drei Schuppen am oberen Rand des Kiemendeckels. Frisch gefangene Tiere sind insgesamt orangerot, mit unregelmäßig verteilten schwach gelben Flecken. Bisher ist die Art nur von Indonesien bekannt, von der Gegend der Inseln Maumere, Flores und Kawe im Raja-Ampat-Archipel. Trimma xanthochrum n. sp. ist durch folgende Merkmale gekennzeichnet: breite Interorbitalregion (80-100% des Pupillendurchmessers), einen zweiten Rückenstachel, der normalerweise nach hinten bis zwischen die Ansätze des zweiten und dritten Rückenflossenstrahls reicht, Brustflossenstrahlen, in der Regel mit 7-8 verzweigten Strahlen, senkrechte Reihen von Sinnespapillen unterhalb vom Auge mit jeweils 2-3 Papillen in den Reihen 1 bis 4 und 4-5 Papillen in Reihe 5, einen Schwanzfleck, dessen untere Hälfte etwa zwei Drittel der Breite der oberen Hälfte einnimmt, sowie normalerweise eine gelbliche Gesamtfärbung auf dem Rumpf, die sich mindestens proximal in einem Gelbton auf der Schwanzflosse fortsetzt. Gegenwärtig kennt man diese Art mit Sicherheit nur vom Raja-Ampat-Archipel. Résumé Une 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 heure connues exclusivement sur ces îles, tandis que les autres peuvent être trouvées dans d autres parties du Pacifique Ouest. Trimma cheni n. sp. se distingue des autres espèces de Trimma par 8-9 écailles sur la ligne médiane prédorsale, 2-3 écailles sur l opercule, un deuxième rayon épineux étiré sur la première nageoire dorsale allant à l arrière jusqu au-dessus des bases des rayons 1-4 de la seconde dorsale, des rayons de la nageoire pectorale divisés au milieu, le cinquième rayon de la pelvienne divisé une fois, une rayure sombre à la base des nageoires dorsales, l attache des écailles indistinctement marquée d une pigmentation plus sombre, et à l état vivant, deux barres rouges à orange sur la joue et trois raies jaunes diffuses sur le corps (les plus apparentes le long du pédoncule caudal). Elle a été recensée aux Palaos, aux Philippines, et au Sulawesi et à Flores en Indonésie. Trimma erdmanni n. sp. présente un corps orange rougeâtre, avec une rayure latérale rouge sombre, aux bordures plus foncées, qui remonte jusqu à la tête, où elle se divise ; là, une fine rayure longitudinale de couleur claire se trouve sous l œil. Elle ne possède habituellement pas d écailles sur la ligne médiane prédorsale, la seconde épine de la première dorsale est étirée, la dorsale compte 9 rayons et l anale 8, une seule division sur le cinquième rayon de la pelaqua vol. 17 no July

3 Richard Winterbottom vienne et branchiospines sur le premier arc branchial. L espèce est connue sur les Îles Raja Ampat, les Îles Hermit et à Madang (Papouasie-Nouvelle-Guinée), et sur des photographies prises à El Nido (Île de Palawan) et dans le Golf de Davao (Mindanoro, Philippines). Trimma habrum n. sp. possède un espacement interorbital osseux égal au diamètre de sa pupille, 8-9 écailles sur la ligne médiane prédorsale, une nageoire pectorale à 14 rayons non divisés, une nageoire pelvienne dont le cinquième rayon n est pas divisé, une absence de membrane basale reliant le cinquième rayon de part et d autre de la ligne médiane, habituellement une seule rangée complète de nageoires sur les joues et des écailles sur les deux tiers supérieurs de l opercule. Les tons frais de sa coloration sont diagnostiques : un dorsum pâle et translucide, avec des taches jaune clair et la base de chaque élément de la dorsale entourée d un point rouge, une barre rouge fine le long des bords postérieurs des hypuraux, une absence totale de pigmentation foncée sur la région hypurale du pédoncule, et une pigmentation sombre couvrant les bordures dorsales de la vessie natatoire, la gaine des neurones et le cerveau. Elle est pour l instant uniquement connue sur le résultat d une seule collecte réalisée sur l Île Kerou (Îles Fam, Raja Ampat, Indonésie). Trimma haimassum n. sp. se caractérise par un corps relativement épais, de nombreuses écailles irrégulières avec environ 30 rangées latérales et plus de 15 écailles dans les séries transversales antérieures, un cinquième rayon de la pelvienne se ramifiant deux fois de manière dichotomique, habituellement une seconde épine dorsale quelque peu étirée, une absence de prédorsale, des écailles sur les joues ou les opercules, un espacement osseux interorbital limité avec une excroissance charnue médiane entre les deux yeux, et une excroissance dermique avant la première épine dorsale. Un point rouge ou une tache étirée au-dessus de l opercule est visible sur des spécimens vivants et frais, et la surface dorsale du museau présente un motif réticulé foncé, avec des rayures sombres transversales sur la bordure dorsale de l orbite. L espèce est connue du nord du Sulawesi sud-occidental jusqu à Palawan et l est des Îles Salomon. Trimma papayum n. sp. est unique parmi les espèces décrites dans le genre pour avoir un point noir ocellé sur la pupille, d un tiers du diamètre de celleci et un autre juste derrière le quatrième rayon épineux de la dorsale. Elle possède une nageoire dorsale à 9 rayons et une anale à 8, un seul point de ramification sur le cinquième rayon de la pelvienne, rangées d écailles transversales antérieures et 8-9 postérieures, 5-10 écailles sur la ligne médiane prédorsale, et une seule rangée de 3 écailles sur la bordure supérieure de l opercule. Les spécimens fraîchement collectés ont une coloration générale rouge orange, ponctuée de points jaunes diffus et éparpillés. L espèce est seulement connue en Indonésie, à Maumere (Flores) et sur l Île Kawe (Raja Ampat). Trimma xanthochrum n. sp. est caractérisée par une large région interorbitale (80-100% du diamètre de la pupille), une deuxième épine dorsale arrivant habituellement, à l arrière, entre les bases du deuxième et du troisième rayon de la dorsale, rayons sur la pectorale dont habituellement 7-8 sont divisés, des rangées verticales de papilles sensitives 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 coloration du corps globalement jaunâtre, avec au minimum du jaune à proximité de la nageoire caudale. Pour l instant, on ne l a identifiée avec certitude que sur les Îles Raja Ampat. Sommario Recenti spedizioni nelle Isole Raja Ampat, Indonesia (2010) hanno portato al campionamento di sette specie non descritte di Trimma, sei delle quali sono descritte in questo articolo. Due di queste sono al momento note solo in queste isole, le altre anche da altre parti del Pacifico occidentale. Trimma cheni n. sp. si distingue dalle altre specie di Trimma per avere 8-9 scaglie sulla linea mediana predorsale, 2-3 scaglie sull opercolo, la seconda spina dorsale allungata fino a raggiungere posteriormente la base dei raggi 1-4 della seconda pinna dorsale, raggi pettorali mediani ramificati, quinto raggio pelvico con una singola ramificazione, una stria basale scura sulle pinne dorsali, avvallamenti delle scaglie indistintamente delimitate da pigmento più scuro e, in vita, due barre rossoarancio lungo le guance e sul corpo tre strie gialle diffuse (più evidenti lungo il peduncolo caudale). È stata rinvenuta a Palau, Filippine e a Sulawesi e Flores in Indonesia. Trimma erdmanni n. sp. ha il corpo arancio-rossastro con una stria laterale rosso scuro con bordi più scuri sul corpo che si estendono anteriormente sul capo, do ve si biforca, ed è presente una sottile striatura longitudinale chiara sotto l occhio. In genere non possiede scaglie predor sa li, ha la seconda spina della prima dorsale allungata, pos siede 9 raggi dorsali e 8 anali, una singola ramificazione nel quinto raggio pelvico e rastrelli branchiali sul primo ar co branchiale. La specie è stata raccolta alle isole Raja Ampat e a Madang e alle isole Hermit (Papua Nuova Guinea) ed è stata fotografata a El Nido, isola di Palawan e nel golfo di Davao, Mindanoro, Filippine. Trimma habrum n. sp. ha una distanza interorbitale ossea uguale al diametro della pupilla, 8-9 scaglie predorsali, 14 raggi pettorali non ramificati, quinto raggio pelvico non ramificato, membrana basale che unisce il quinto pelvico alla linea mediana assente, in genere una singola fila completa di scaglie sulla guancia e scaglie sui due-terzi superiori dell opercolo. La colorazione dell esemplare fresco è diagnostica: il dorso è traslucente e pallido con macchie giallo chiaro e la base di ogni elemento della dorsale circondato da una macchia rossa, una sottile barra rossa lungo i margini posteriori degli ipurali, nessuna pigmentazione scura nella regione ipurale del peduncolo e una pigmentazione scura che ricopre i margini dorsali della vescica natatoria, la guaina neurale e il cervello. Al momento è nota solo da un singolo campionamento eseguito a Kerou Island, Isole Fam, Raja Ampat, Indonesia. Trimma haimassum n. sp. si caratterizza per avere un corpo relativamente alto, numerose scaglie irregolari presenti in 30 file laterali più di 15 scaglie in linea anteriore trasversale, il quinto raggio pelvico che si ramifica due volte in modo dicotomico, la se conda spina dorsale generalmente allungata, assenza di scaglie predorsali e anche sulla guancia e sull opercolo, l interorbitale moderatamente largo con una cresta mediana carnosa tra gli occhi e una cresta dermica anteriore alla prima spina dorsale. È presente una macchia rossa o una chiaz za allungata sopra l opercolo negli esemplari appena 129 aqua vol. 17 no July 2011

4 Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature METHODS Methods and the format of the descriptions follow Winterbottom (2002), except that pectoral and pelvic-fin ray branching is described from preserved material stained with a cyanine blue solution as outlined in Akihito et al. (1993; Akihito et al. 2002: 1270). Lengths given are Standard Length (SL) in millimetres; SD = Standard Deviation; values for the holotypes are given in bold where appropriate. In the additional (non-type) material examined sections, the catalogue number is followed by the number of specimens in the lot (except where there is only a single specimen) with the range of SL given in parentheses. Abbreviations for repositories of material examined follow the codon abbreviations published by the American Society of Ichthyologists and Herpetologists ( The letters CS after a ROM catalogue number denote specimens that were cleared with trypsin and counter-stained with alizarin and alcian blue. The vertebral transition type (A or B) between the abdominal and caudal vertebral centra is defined and illustrated by Winterbottom (2003: Fig. 14 (inset)), and Winterbottom & Zur (2007: Fig. 3). Essentially, in type A, the last two abdominal vertebrae are of the usual perciform configuration in that they lack a bony connection across the midline between the bases of the paired haemal arches, and the first (and often subsequent two to three) caudal vertebrae have a single greatly enlarged canal with only the tips of the haemal arches fused together in the midline. In type B, the last two abdominal vertebrae have a bridge joining the haemal arches across the midline, forming a haemal canal; the first caudal vertebra has broad haemal arches with a small basal haemal canal and a larger secondary canal distal to this, which forms a posteriorly tapering funnel and embraces the posterior end of the swimbladder. Counts and measurements were input directly into an Excel file with Mitutoyo digital callipers using WinWedge 3.01 software. Photographs other than the portraits of fresh or live specimens were produced from multiple digital images taken with a Nikon D100 or D300S camraccolti, mentre la superficie dorsale del muso mostra un motivo reticolato scuro, con striature trasversali scure sopra il margine dorsale dell orbita. La specie è nota dal Sulawesi sudoccidentale a nord fino a Palawan e a est fino alle Isole Salomone. Trimma papayum n. sp. è unica tra le specie de - scritte in questo genere per avere una macchia ocellata nera dal diametro pari a un terzo a quello della pupilla sopra e ap - pena dietro la quarta spina dorsale. Possiede 9 raggi dorsali e 8 anali, un singolo punto di ramificazione nel quinto raggio pelvico, file di scaglie trasversali anteriori e 8-9 po - steriori, 5-10 scaglie predorsali e una sola fila di 3 scaglie sul mar gine superiore dell opercolo. Esemplari appena raccolti hanno una livrea nel complesso rosso-arancio, con mac chie gialle diffusamente disseminate. La specie è nota solo in Indonesia, a Maumere, Flores e Kawe Island, Raja Ampat. Trimma xanthochrum n. sp. è caratterizzata da un ampia regione interorbitale (80-100% del diametro della pupilla), da una seconda spina dorsale che raggiunge di solito posteriormente lo spazio tra le basi del secondo e il terzo raggio dorsale, raggi pettorali con generalmente 7-8 raggi ramificati, file verticali di papille sensoriali sotto l occhio formate da 2-3 papille nelle file 1-4 e 4-5 nella fila 5, una chiazza caudale che ha la metà inferiore circa due terzi l ampiezza della metà superiore e, infine, una colorazione generalmente giallastra con del giallo almeno prossimalmente nella pinna caudale. Attualmente è nota con certezza solo alle Isole Raja Ampat. INTRODUCTION Trimma Jordan & Seale (type species: T. caesiura Jordan & Seale, 1906) contains about 100 species of small (<30 mm SL), often colourful gobiids, primarily associated with Indo-Pacific coral reefs. Members of the genus may be recognized by the lack of cephalic sensory canal pores, a much reduced cephalic sensory papillae pattern, a wide gill opening extending anteriorly to below the vertical limb of the preopercle or, more usually, anterior to this, a lack of spicules (odontoids) on the outer gill rakers of the first gill arch, fewer than 12 dorsal and anal fin rays, and a fifth pelvic-fin ray that is equal to or more than 40% the length of the fourth pelvic-fin ray (Winterbottom 1995, 2003). Although the gill rakers on the first gill arch of members of this genus often have a row of crenulations or serrations (a series of sharply rounded protrusions lacking any ossification) along their medial margins (Fig. 1), these structures are not considered here as the equivalent of odontoids. There are currently 64 valid species of Trimma, with approximately 35 additional known undescribed species (Winterbottom & Hoese, unpublished). A recent (January/February, 2010) rapid survey of the Raja Ampat islands off the Bird s Head region of New Guinea sponsored by Conservation International s Indonesia Marine Program resulted in the collection of seven undescribed species of this genus. aqua vol. 17 no July 2011 This paper describes six of those new species. Additional specimens of some of these new species were obtained from the Raja Ampat area by Mark Erdmann on subsequent visits. The species described here were listed under their RW numbers by Dimara et al. (2010). 130

5 Richard Winterbottom Table I. Counts of cephalic sensory papillae in the different rows of the six new species of Trimma. Values consist of minimum, 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) (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) (11.9, 1.48, 20) 9-11 (9.9, 0.7, 10) 6-10 (7.6, 1.07, 20) Row e ant (13.5, 1.63, 19) (16.5, 2.11, 19) (11.8, 1.28, 12) (19.1, 1.89, 20) (16.2, 1.08, 10) 12-19(14.5, 1.67, 19) Row e pos (14.1, 1.71, 19) (19.2, 1.85, 19) (13.3, 1.48, 11) (20.7, 2.31, 20) (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 (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 (13.8, 1.6, 10) (17.6, 1.38, 18) (12.8, 1.03, 9) (18.1, 1.24, 20) (16.1, 1.22, 10) (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) era attached to a Zeiss SV-8 dissecting microscope at slightly incremental focal planes selected manually, or with a Canon EOS Rebel XS camera attached to a Zeiss SV-12 dissecting microscope using Zeiss AxioVision 4.8 software and automatic increments. The images were then collated into a single image using Helicon Focus 5.1 (HeliconSoft). All images, drawings and image collations were made by the author unless otherwise noted. Type specimens are confined to those collected 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 surface of the cheek. Sanzo (1911) uses this letter for both the row immediately behind, and which fol- lows the orientation of, the maxilla and for a horizontal row across the lower cheek that is, at least in Trimma, well separated from the first row anteriorly. Miller (1972a) refined this nomenclature, using d for the row behind the maxilla, and reserving d for the row across the cheek, and this distinction is followed in this paper. [Note: Miller (1972a; 1972b) actually uses d and d 1 respectively in the text of both papers and for the figure in the 1972b paper, but employs d in Figure 5 of the 1972a publication. Since Sanzo (1911) consistently uses prime where appropriate for other rows of papillae, I have opted to follow the nomenclature used in Miller s figure (1972a, Fig. 5)]. This separation of row d into two discrete entities is also found in both Priolepis and Egglestonichthys (Miller 131 aqua vol. 17 no July 2011

6 Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature & Wongrat 1979; Winterbottom & Burridge 1992). It is also present in the possibly related Lythrypnus (Ahnelt & Bohacek (2004) - where the authors referred to the two sections as d 1 and d 2 for the anterior and posterior sections respectively). However, such a separation is apparently rare in gobies (D. F. Hoese, in litt.). The complete nomenclature of the papillae rows used here is given in Figure 2A. Sanzo s (1911) nomenclature was largely accepted and used by Aurich (1938), although a brief comparison of the two works brings several apparent discrepancies to light. Hoese (1983) pointed out some of the difficulties in homologizing Sanzo s (1911) names with the papillae patterns he found in Indo-Pacific gobies, and proposed a revised nomenclature based on whether the bases of the papillae were longitudinally or vertically oriented with respect to the axis of the papilla row. Takagi (1988) also proposed a Fig. 1A-C. Left lateral view of the upper portions of the first gill arches to show crenulated edges of gill rakers in: A. Trimma erdmanni (26.2 mm SL paratype, ROM 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, C cleared and stained with alizarin and alcian blue note lack of crenulations, indicating that they are structures without any calcification. aqua vol. 17 no July 2011 Fig. 2A-B. Composite image of head to show distribution and 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 grey connecting lines in A represent the rows as counted in this paper, those in B represent parts of the nerves that stained in the cyanine blue preparation. 132

7 Richard Winterbottom system of nomenclature for the papillae rows, and provided a table equating his names with those used by previous authors (Takagi 1988: Tbl. 4). Other authors have been content to simply illustrate the papillae patterns (although Akihito et al. (1988: Fig. 36) did distinguish between the transverse and longitudinal rows). Trimma has been characterized as having a reduced transverse papillae pattern, consisting only of longitudinal rows of papillae on the cheek (Winterbottom & Burridge 1992; Hoese et al. 2011). One of the putative sister groups of Trimma, Priolepis (Winterbottom 1984; Miya, pers.comm.) has several transverse rows of cheek and interorbital papillae, at least in the hypothesized basal members of that clade (Winterbottom & Burridge 1992: Fig. 12). Most species of Trimma have two longitudinal rows of papillae (a and c) on the cheek immediately under the eye and above row d. Frequently, the anteriormost papillae in rows a and c lie below the anterior border of the eye forming a vertical row of two papillae, and the papillae below the posterior margin of the pupil form a vertical row of two or three papillae (in the later case potentially attributable, from dorsal to ventral, to one papilla from each of rows a and c and a single papillae named cp by Sanzo (1911), respectively). Ahnelt & Bohacek (2004) also employ this nomenclature for the ventralmost papilla in Lythrypnus, but Asaoka et al. (2011: Fig. 4) use cp for the entire vertical row, which consists of at least five, perhaps more, individual papillae in Pterogobius elapoides. Ahnelt & Göschel (2003: Fig. 3) also use cp for a vertical row of several papillae on the cheek in Quietula. One of the new species described here (Trimma xanthochrum) has two to five papillae in each of five vertical rows in this position. Where such vertical rows occur, Sanzo (1911) employs numbers 1 through 6 as identifiers, beginning with 1 for the row below the anteroventral margin of the eye. The fifth row, below the posterior margin of the pupil, usually marks the anterior limit of row b. Given the hypothesis that Trimma exhibits a reduced transverse pattern of cheek papillae (with T. xanthochrum possessing the most plesiomorphic condition in the genus), the application of Sanzo s (1911) nomenclature of rows a and c for the longitudinal papillae rows below the eye is incorrect since these papillae represent the remnants of the vertical rows 1-5 (see also Hoese et al. 2011). An alternative nomenclature is given in Figure 2B, based on an interpretation that the papillae rows below the eye in Trimma do represent originally more extensive vertical rows of papil- lae. It seems probable that when homologies can be empirically established, a single, unified (and different) nomenclature for papillae will need to be adopted. The recent explorations of the innervations of the different papillae patterns in various gobioids (e.g. Ahnelt & Bohacek 2004; Asaoka et al. 2011) is a promising approach, but also raises some of the difficulties envisioned by Hoese (1983). For example, the phylogenetic homology of the papillae in the rows a, c and cp may prove especially difficult to resolve. Most of the head papillae have an ovoid base, and the long axis is nearly always oriented at right angles to the path of the nerve supplying them. Apparent exceptions to this in most species of Trimma, such as rows a, c and p, where the path of the nerve is aligned with the long axis of the bases of the papillae, may well prove to be the result of reduction to a single papilla of originally transverse rows that had the papillae bases transversely oriented with respect to their immediate nerve track. Since the papillae patterns in Trimma correspond well, for the most part, with those illustrated and labelled by Sanzo (1911: Pl. 9, Figs 9-10) for Gobius affinis Kolombatovic, 1891 (current status: Pomatoschistus pictus adriaticus Miller, 1973), Sanzo s nomenclature is employed here with the exception noted above (Fig. 2A). Although the identity of all the rows of papillae labelled in figures of Trimma in this paper appear to be positional homologues of Sanzo s rows, they should not be construed as phylogenetic homologues. Note also that the grey line linking individual papillae into the named rows as used in this paper (Fig. 2A) is not based on an examination of the innervations or of the phylogenetic history of the genus, and it is probable that some papillae are incorrectly attributed to or omitted from that row. The papillae are easily abraded from the surface of the head (especially those in rows b, d, z and g), and are usually difficult to see where the area is covered with scales. Where counts are given, they should thus be regarded as minimum values. Those counts exhibiting large variation probably do so because 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) 133 aqua vol. 17 no July 2011

8 Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature Material Examined: A total of 8 lots, 56 type specimens ( mm SL), plus an additional non-type specimen (tissue voucher), all from the Raja Ampat Islands, Indonesia, and other specimens from Indonesia and the Philippines. The description is based on the holotype and 19 other specimens from ROM 85161, 85138, and 1835CS, mm SL (x = 19.3, SD = 1.31), 12 males and 8 females. Holotype: ROM 87481, 20.0 mm SL male, Tanjung Manare, Waigeo Island, west side at about middle of width off small cape ( S, E), 52 m, rotenone, , field # RW10-24, 29 January 2010, M. V. Erdmann. Paratypes: AMS I (formerly ROM 85221), 3, , Wofoh Island, west coast near south end ( S, E), 48 m, clove oil, , field # RW10-26, 29 January 2010, M. V. Erdmann. MZB (formerly ROM 87479), 10, , Waigeo Island, Tanjung Manare ( S, E), 52 m, clove oil, 28 May, 2010, M. V. Erdmann. ROM 85085, 4, , about 1.2 km SSE of Mutus Island, on west side of sand spit ( S, E), 50 m, clove oil, , field # RW10-16, 28 January 2010, M. V. Erdmann. ROM 85138, 11, , Jef Tsiep Island, west side, channel between it and small island to the west ( S, E), m, clove oil, , field # RW10-20, 28 January 2010, R. Winterbottom, L. Katz & P. Johannes. ROM 85161, 17, , Tanjung Manare, Waigeo Island, west side at about middle of width off a small cape ( S, E), m, rotenone, , field # RW10-23, 29 January 2010, R. Win- 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. aqua vol. 17 no July

9 Richard Winterbottom terbottom, R, L. Katz, P. Johannes, W. Kaka & W. Awom. ROM 85197, 7, , collected with the holotype. ROM 85315, 1, 17.6, S of Misool Island, off Waaf Island ( S, E), clove oil, , field # RW10-36, 1 February 2010, M. V. Erdmann. ROM 1835CS (formerly ROM 85232), 2, , Wofoh Island, west coast near south end ( S, E), m, clove oil, , field # RW10-27, 29 January 2010, L. Katz. Additional (Non-type) Material. Raja Ampat: ROM T07737, (21.8), collected with ROM (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); ROM 53114, 3 (20-21). Negros Oriental, ROM 53112, 9 (12-22); ROM 53113, (18). Siquijor, ROM 53107, (19); ROM 53108, 6 (19-22); ROM 53109, 8 (16-21); ROM 53110, 11 (10-22); ROM 53111, 8 (21-24). Diagnosis: Trimma cheni has 8-9 scales in the predorsal midline, a row of 2-3 usually ctenoid scales along the upper border of the opercle, no cheek scales, a dark basal stripe in the dorsal fins, an elongate second dorsal spine reaching posteriorly to the bases of rays 1-4 of the second dorsal fin, the middle rays of the pectoral fin branched, a fifth pelvic fin ray which branches once dichotomously, the basal membrane connecting the inner margins of the fifth pelvic fins rays is less than 15% the length of the fifth ray, the scale pockets are not distinctly outlined with darker pigment, and, in life, there are two red to orange bars across the cheek 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- teriorly to bases of rays of second dorsal fin when adpressed (x = 2.3, SD = 1.02, n = 19), rays all branched except, usually, posterior element of last ray (first ray unbranched in 2), last ray reaching posteriorly 40-50% of distance from its base to first dorsal procurrent ray (x = 48.8, SD = 3.05); anal fin I 8, all but first and posterior element of posteriormost ray branched; posteriormost ray % of distance from its base to first ventral procurrent ray (x = 41.0, SD = 2.55, n = 19); pectoral fin (x = 17.1, SD = 0.38), (mean = 5.4, SD = 1.03) and 4-8 (x = 5.7, SD = 1.36) unbranched dorsal and ventral rays respectively, middle rays branched, fin reaching posteriorly to a vertical line between bases of anal rays 1-3 (x = 1.3, SD = 0.56, n = 19); pelvic fin I 5, no frenum, basal membrane reduced and % length of fifth ray (x = 13.0, SD = 1.91), first four rays with one sequential branch, fifth ray branched once and % length of fourth (x = 52.6, SD = 2.50, n = 18), fourth ray reaching posteriorly to between bases of anal rays (x = 4.0, SD = 1.24, n = 17). Lateral scales 23; anterior transverse scales (x = 8.1, SD = 0.37); posterior transverse scales (x = 7.5, SD = 0.41); predorsal scales 8-9 (x = 8.2, SD = 0.39); no scales on cheek; opercle with single row of 2-3 usually ctenoid scales; pectoral base with usually 3 vertical rows of scales and 4 cycloid scales in posterior row; (x = 6.4, SD = 0.79) cycloid prepelvic scales (in midline anterior to basal membrane); (x = 11.9, SD = 0.3) circumpeduncular scales; body scales ctenoid except for cycloid scales on anterior belly midline and on, beneath and just posterior to pectoral fin base; body scales extending anteriorly to just behind eye. Gill opening extending anteroventrally to a vertical below anterior onethird of pupil. Upper jaw with outer row of curved, enlarged, spaced canines along length of premax- Fig. 5. Left lateral view of Trimma cheni, 17.6 mm SL juvenile paratype, ROM 85315, Waaf Island, Raja Ampat. Note the parasitic copepod attached immediately posterior to pelvic fins. Photo by R. Winterbottom. 135 aqua vol. 17 no July 2011

10 Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature illa, gradually decreasing in size posteriorly until not much larger than inner rows; two to three irregular inner rows of smaller teeth almost reaching distal tip of premaxilla, innermost row of slender, slightly curved, spaced teeth almost as long as outer row, from symphysis to bend of jaw in males and some females. Lower jaw with outer row of 4-5 curved, enlarged, spaced canines reaching to bend of dentary, several irregular inner rows of slightly curved caniform teeth half size of outer, becoming reduced in size and less curved posteriorly and tapering to form single row of straight conical teeth at beginning of coronoid process; innermost row of enlarged slightly curved teeth at symphysis, increasing in size along jaw (equal in size to outermost row just behind bend of dentary, then gradually decreasing in size to dorsal tip of coronoid process (becoming equal in size to teeth in inner rows). Cephalic sensory papillae row counts given in Table I. Tongue truncate with rounded edges. Gill rakers on first arch = (x = 19.2, SD = 0.59). Anterior nasal opening a short tube extending out over upper lip, posterior nasal opening a pore with raised rim, both protruding from slightly raised oval sac confined to anterior half of snout. Bony interorbital % (x = 38.1, SD = 4.08) pupil width, with broad moderately developed U- shaped interorbital furrow and shallow postorbital groove (or none). No ridge of tissue (or dermal crest) extending anterior to first dorsal spine. Epaxialis musculature extending anteriorly to above vertical with posterior margin of pupil. Last two abdominal vertebrae with bridge of bone connecting haemal arches just distal to their bases, forming haemal canal bounded dorsally by centrum. First caudal vertebra with two canals, a small proximal one at base of haemal arches, which fuse in midline briefly before diverging again to form posteriorly directed funnel-like second arch. Haemal arches then fusing again in midline to form haemal spine (vertebral transition Type B). Colour pattern (from images of four freshly collected specimens, mm SL). A 22.3 mm SL male (Fig. 3) has a mostly red head, darker on the snout, with two lighter and more intense red bars with irregular diffuse yellowish margins on the cheek below the anterior and middle of the eye, and another yellow-brown bar over the vertical limb of the preopercle, the dorsal half of the opercle with a similar colouration; the posterior margin Fig. 6. Map showing the distributions of the new species of Trimma described. all species present; T. cheni; T. erdmanni; T. haimassum; T. papayum; T. xanthochrum. aqua vol. 17 no July

11 Richard Winterbottom of the branchiostegal membrane immediately adjacent to the opercle and subopercle is orange. The iris is red grading to yellow orange ventrally, with an anteroventral to posterodorsal dark purple stripe across the pupil; the dorsal surface of the orbit has several alternating red and white stripes passing across the midline. The body is yellowish, grading to pink on the peduncle, with three vague yellow stripes (most obvious on the peduncle), with heavily scattered dark chromatophores, especially on the dorsum and along the scale pockets and scale margins. The membranes of the dorsal fins are densely speckled with melanophores, except for a half-pupil diameter wide yellow stripe just above the base and, above this, a second but more diffuse yellow stripe. The membranes of the caudal fin are similarly invested with melanophores, with several streaks and blotches of yellow separated by three darker diffuse stripes. The distal margin of the anal fin is dusky, and there is a broad (about one-and-ahalf pupil diameters wide) yellow stripe with diffuse margins and a dark basal stripe. The pectoral and pelvic fin rays are yellow or orange, with hyaline membranes, the bases of the fins are light tan with many scattered dark brown chromatophores. A 20.1 mm SL male is similar, but with a darker and greyer body, while a 20.0 mm SL male is paler, with a generally more yellow cast. A specimen from El Nido, Philippines photographed underwater (Fig. 4) has a greyish background with diffuse yellow-green markings on the body, three bars on the lower part of the head (two below the eye, the third over the vertical limb of the preopercle), which grade from orange yellow dorsally into the greenish-yellow ventrally in the region of the lower jaw and branchiostegals. There are two short oblique orange bars passing posterodorsally from the posterior margin of the upper orbit, and three short orange to red transverse bars across the dorsal margin of the eye. Other specimens photographed underwater have a redder head and either a greenish-yellow or brownish body. A 17.6 mm SL juvenile (Fig. 5) has a relatively dark body, two dark red bars on a grey-pink cheek, and the stripes in the fins are reddish rather than yellow. Colour pattern in alcohol: strawcoloured with a heavy scattering of brown chromatophores and melanophores on the body, nape and pectoral fin base. The opercle has numerous densely scattered melanophores. The cheek is paler, and two diffuse lighter bars (corresponding to the red bars in fresh material) may or may not be appar- ent. The dark basal stripes in the dorsal and anal fins remain obvious, the dark stripe at the midregion of the second dorsal fin is relatively well defined, and three diffuse dark stripes run parallel to the rays of the caudal fin. The dorsal rim of the pectoral fin base is often somewhat darker than the surroundings, appearing as a diffuse dark streak. Comparisons: Trimma cheni belongs to a group of species which has a scaled predorsal midline, no cheek scales, a scaled opercle, at least the middle rays of the pectoral fin are branched, and a dark basal stripe in the dorsal fins is present (Winterbottom 2006). Within this group, T. cheni appears closest morphologically to T. milta Winterbottom, 2002, in having a poorly developed postorbital trough or none and in possessing a single row of 2-3 scales on the opercle. However, T. milta does not have the second dorsal spine elongated, has an unbranched fifth pelvic fin ray, has the scale pockets on the body lightly outlined with darker pigment, and lacks the conspicuous red to orange bars on the cheek as well as the three yellow stripes on the body. Trimma anthrenum Winterbottom, 2006, differs in lacking opercular scales and elongate second dorsal spine, in possessing a well developed postorbital trough, and in live colouration (plain yellow with a distinctively coloured iris (see Winterbottom 2006: Figs 1-2). Trimma preclarum Winterbottom 2006 differs from T. cheni in lacking scales on the opercle, in possessing a well developed postorbital trough, and in lacking the two red to orange bars on the cheek, although this species does have three yellow stripes on the posterior part of the body. These stripes are also evident in T. hayashii Hagiwara & Winterbottom, 2007, which lacks scales in the predorsal midline (or, if scales are present, they are represented one or two scales crossing the midline between the eye and the first dorsal fin origin) and on the opercle, and has a distinctive black ocellated spot on the branchiostegal membranes below the vertical limb of the preopercle. Distribution: Trimma cheni is currently known from the Raja Ampat islands, and from Palau, the Philippines and south-west to Sulawesi and Flores in Indonesia (Fig. 6) at depths of 5-52 m. Etymology: The species is named for I.-S. Chen, National Taiwan Ocean University, Keelung, Taiwan, in acknowledgment of his numerous publications on Indo-Pacific reef fishes in general, and gobies in particular. This species has been referred to as Trimma RW sp aqua vol. 17 no July 2011

12 Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature 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 Peninsula) Trimma RW sp. 68 Dimara et al. 2010: 621 (Raja Ampat) Trimma sp. Allen et al. 2003: 329 (lower right), Philippines Material Examined: A total of 7 lots, 64 type specimens, plus four additional non-type specimens (tissue vouchers), and three additional lots of non-type specimens from Papua New Guinea. All type specimens collected at the Raja Ampat Islands, Indonesia. Description based on up to 24 specimens from ROM 85033, 85144, 85186, 87460, and 1837CS ( mm SL, 12 males, 12 females). Holotype: ROM 87482, 23.5 mm SL male, Fig. 7A-B. Head of Trimma erdmanni (19.2 mm SL female paratype, ROM 85033) in: A. left lateral and B. dorsal view. Specimen stained with cyanine blue. Photo by R. Winterbottom. aqua vol. 17 no July 2011 Kawe Island, SW bay ( S, E), 50 m, clove oil. M. V. Erdmann, 26 January Paratypes: AMS I (formerly ROM 85330), 1, 23.1, Kepotsol Island, east side ( S, E), 66 m, clove oil, field # RW10-38, , 1 February 2010, M. V. Erdmann. MZB (formerly ROM 87460), 7, , Waigeo Island, Tanjung Manare ( S, E), 52 m, clove oil, no field No., 28 May, 2010, M. V. Erdmann. ROM 85033, 22, , collected with the holotype. ROM 85144, 8, , Jef Tsiep Island, west side, channel between it and small island to the west ( S, E), 42 m, clove oil, field # RW10-21, , 28 January 2010, M. V. Erdmann. ROM 85186, 16, , Tanjung Manare, Waigeo Island, west side at about middle of width off small cape, sponges, hard corals ( S, E), 52 m, 1 kg powdered rotenone & detergent, field # RW10-24, , 29 January 2010, M. V. Erdmann. ROM 85393, 7, , SE islands off Misool, south side of Balbulol Island ( S, E), 45 m, 1 kg powdered rotenone & detergent, field # RW10-45, , 2 February 2010, M. V. Erdmann. ROM 1837CS, 4, , Wofoh Island, west coast near south end ( S, E), 48 m, clove oil, field # RW10-26, , 29 January 2010, M. V. Erdmann. Additional (Non-type) Material. Raja Ampat: ROM T07723 and T07724, 2 ( ), collected with ROM 85033; ROM T07743 and T07744, 2 ( ), collected with ROM Papua New Guinea. Hermit Islands: USNM , 17 ( Madang: WAM P , (19.0); WAM P , (21.2). Diagnosis: Trimma erdmanni differs most trenchantly from other species of the genus in the colour pattern of a red to orange body with a darker red to orange lateral stripe with dark borders on the body which extends anteriorly onto the head, where it bifurcates, and there is a thin light stripe from the maxilla to the opercle that abuts the ventral margin of the eye. The new species usually lacks scales in the predorsal midline, but when these are present, they are separated from the first dorsal fin by an unscaled area. The second (and third to some degree) spines of the first dorsal are elongated, there are 9 dorsal and 8 anal rays, the middle rays of the pectoral fin are branched, there 138

13 Richard Winterbottom is a single branch in the fifth pelvic fin ray, the basal membrane is 7-18% of the length of the fifth fin ray and there are gill rakers on the outer margin of the first gill arch. Description: Dorsal fins VI + I 9, second and third spines longest, second spine reaching to between bases of second to eighth rays of second dorsal fin, third spine reaching to between bases of dorsal spine and fourth ray of second dorsal fin when adpressed, first ray of second dorsal fin usually unbranched (branched in 5, SD = 0.41), posterior element of last ray unbranched, last ray extending between % (x = 74.2, SD = 16.0) of distance between its base and first dorsal procurrent caudal fin ray; anal fin I 8 (n = 24), all but first and posterior element of posteriormost ray branched, last ray extending between 40-80% (x = 60.7, SD = 12.4) of distance between its base and first ventral procurrent caudal fin ray; pectoral fin (x = 18.0, SD = 0.61), 3-4 (x = 3.4, SD = 0.49) and (x = 3.4, SD = 1.00) unbranched dorsal and ventral rays respectively, middle rays branched, fin reaching posteriorly to a vertical line between anterior margin of urogenital papilla and base of first ray of anal fin; pelvic fin I 5, no frenum, basal membrane poorly developed, Fig. 8. Left lateral view of Trimma erdmanni, 26.2 mm SL male paratype, ROM 85033, Kawe Island, Raja Ampat. Note that the 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. 139 aqua vol. 17 no July 2011

14 Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature 18% length of fifth pelvic ray), first four rays with one or two sequential branch points, fifth ray branched once and % length of fourth (x = 64.5, SD = 4.44, n = 20), fourth ray reaching posteriorly to between anterior margin of urogen- tial papilla and base of first anal-fin ray. Lateral scales (x = 23.1, SD = 0.30, n = 20), anterior 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 Fig. 10. Left lateral view of Trimma erdmanni, 23.5 mm SL male holotype, ROM 87482, Kawe Island, Raja Ampat. Photo by R. Winterbottom. Fig. 11. Trimma erdmanni, 14.3 mm SL juvenile paratype, ROM 85033, Kawe Island, Raja Ampat. Photo by R. Winterbottom. Fig. 12. Left lateral view of Trimma erdmanni, 10.2 mm SL juvenile paratype, ROM 85033, Kawe Island, Raja Ampat. Photo by R. Winterbottom. aqua vol. 17 no July

15 Richard Winterbottom in two specimens with one or two cycloid scales respectively on upper part of opercle); midline of predorsal usually scaleless, but with 6 scales in holotype, 4 in a 20.9 mm SL female, and 6 scales in front of first dorsal spine followed anteriorly by a naked area and then 2 more scales across midline in a 26.2 mm SL male; usually 4 cycloid scales in posterior vertical row on pectoral base (once 5), with two vertical rows of scales anterior to this; (x = 7.2, SD = 0.73, n = 20) prepelvic cycloid scales in midline anterior to pelvic-fin base; 12 circumpeduncular scales; 8-9 (x = 8.1, SD = 0.31, n = 18) in midline between base of last anal ray and first procurrent caudal ray; body scales ctenoid except for cycloid scales on anterior belly midline, beneath and immediately posterior to pectoral-fin base, anterior scales on sides of nape, and along base of first dorsal fin; generally, body scales extend anteriorly on sides of nape to between a vertical above anterior margin of opercle and just posterior to posterior margin of eye (Fig. 7). Gill opening extending anteroventrally to a vertical below pupil. Upper jaw with outer row of curved, spaced, enlarged canines, decreasing in size posteriorly to distal tip of premaxilla, a band of several irregular rows of small conical teeth at symphysis, number of rows decreasing posterolaterally and ending just posterior to bend of dentary, innermost row slightly larger and directed posteriorly at symphysis, decreasing in size and becoming vertically oriented, and extending to distal tip of premaxilla. Lower jaw with an outer row of curved, spaced, enlarged canines ending at bend of dentary, several rows of small, slightly curved conical teeth at symphysis, grading to a single row at bend of dentary and continuing to lower coronoid process as a single outer row, innermost row at bend of dentary abruptly larger than other inner teeth (twice the height) in males, decreasing somewhat in size posteriorly and ending at the upper margin of the coronoid process. Cephalic sensory papillae as in Table I and Fig. 7. Tongue truncate with rounded edges. Gill rakers on first arch = (x = 20.4, SD = 0.73, n = 21), gill rakers with obvious fleshy crenulations (Fig. 1A). Anterior nasal opening a short tube extending out over upper lip, posterior nasal opening a pore with a raised rim, both protruding from slightly raised oval sac confined to anterior half of snout. Bony interorbital % pupil width (x = 42.6, SD = 5.26, n = 20), with broadly U-shaped interorbital trench with sloping sides and no postorbital trench. No dermal crest anterior to first dorsal fin. Epaxialis musculature extending anteriorly to about a vertical with posterior margin of pupil (Fig. 7B). Abdominal/caudal vertebral pattern is Type B, but haemal arch of second caudal vertebra is somewhat enlarged. Colour pattern (from slides of six freshly collected specimens, mm SL). A 26.2 mm SL male (ROM 85033, Fig. 8) has an overall pinkish hue, darker on the dorsum, dominated by a slightly more than pupil-width midlateral orange stripe bordered with red, which continues anteriorly onto the head behind the eye in a shallow > - shape, the lower limb touching the ventral margin of the eye, the upper limb to a point in line with the dorsal margin of the pupil. In this specimen, the two arms of the > are joined across the interspace by a narrow red-orange bridge, the interspace is pinkish-grey. The red-orange colouration continues onto the snout anterior to the orbit and the anterior part of both upper and lower lips. The cheek is pale pinkish grey, bordered dorsally by a thin (about one-fifth pupil-diameter in width) bluish-white stripe which ends at the vertical limb of the preopercle. The pectoral fin base and isthmus are pale grey. A reddish-orange stripe curves posterodorsally from the upper margin of the orbit, ending above the middle of the opercle. The nape is darker red than most of the rest of the body, and the dorsal half of the caudal peduncle is somewhat less so, with the space beneath the dorsal fins pale pink. Most of the scales on the dorsal half of the body have a very thin margin of red chromatophores along their posterior margins. The rows above the lateral scales and the row beneath it have a wider band of darker red pigmentation. The abdomen is pale pink, which grades into pale red posteriorly. The midlateral stripe is flanked by diffuse darker stripes about equal in width, formed by scattered dark chromatophores, which also underlie the red stripe, especially in the abdominal region. The elements of the first dorsal fin each have a yellow spot just distal to the base, and there is a very diffuse, half-pupil diameter dark stripe above the spots, with a scattering of melanophores in the fin membrane above this. The anterior elements of the second dorsal fin have similar yellow spots, and melanophores are densely scattered throughout the fin membrane. The central rays of the caudal fin have splotches of yellow a little distal to their bases, with an overall rosy hue (especially proximally and centrally), the membranes of 141 aqua vol. 17 no July 2011

16 Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature the outer part of the fin appears greyish due to numerous scattered melanophores. The anal fin has a faint pinkish hue proximally with many melanophores scattered in the membranes distally. Pectoral and pelvic fin membranes are hyaline, the rays of the median fins and the pectoral fin are pale red. The iris is golden-yellow, with scattered red pigmentation, and heavily invested with dark pigmentation (especially dorsally), with a thin white inner rim, and a dark blue band which crosses the iris from the anteroventral to the posterodorsal margins. A live specimen (Fig. 9) is generally similar, although the interspace of the > part of the lateral stripe on the head almost totally occluded, and the yellow spots in the dorsal fins are hardly discernable. The snout is orange, with the posterior nasal opening in a small white spot, a similar spot just in fron of the eye, a median white stripe from the anterior interorbital region to the level of the posterior nares and another similar short stripe from the posterior interorbital region over the dorsum. In a 20.3 mm SL female (from the same lot), the mid-part of the lateral stripe is more yellow than orange, and the yellow spots of the dorsal fins form stripes. A 23.5mm male (Fig. 10, holotype) has the whole body more heavily invested with melanophores, especially anteriorly, and the pinks and oranges are replaced by yellow. The interspace of the > on the side of the head is almost occluded by the two arms of the stripe, the yellow spots in the first dorsal fin and anterior part of the second dorsal fin form diffuse stripes, the dark stripe above this is less discernable, but there is a diffuse dark stripe above the yellow in the second dorsal fin; in both fins, there is a much greater concentration of melanophores. The basal region of the anal fin is pale yellow rather than pink. A 14.3 mm SL juvenile is very dark overall, with densely scattered dark chromatophores on the head and body (Fig. 11). The orange-red lateral stripe is only discernable on the head and scarcely at all on the body, and the yellow stripes in the dorsal fins are distinct. A 10.2 mm SL juvenile (Fig. 12) has a translucent body and head except for the dark chromatophores, which are accentuated along the midlateral orange-yellow stripe. The upper lip, snout, top of the head and > arms of the lateral stripe are yellow, and the median fins have a scattering of melanophores. A live specimen photographed in Davao Gulf, Mindanao by Arthur Bos has the posterior region of the lateral red stripe expanded posteroventrally to a blunt triangular aqua vol. 17 no July 2011 wedge over the hypural region and anterior bases of the caudal fin rays. Colour pattern in alcohol. The 26.2 mm SL male is straw coloured, including all areas that are red, orange or yellow in fresh specimens; however, most of the dark chromatophores and the melanophores remain obvious. The centre of the body stripe above the pectoral fin has a diffuse posteriorly tapering darker stripe formed by scattered small chromatophores, and the margins of the main stripe are bordered dorsally and ventrally by a dark diffuse pupil-width stripe of melanophores that tapers somewhat posteriorly and ends at the hypural plate. A few small melanophores are scattered over the body. The V-shaped anterior bifurcation of the stripe is not apparent anteriorly, but each arm of the bifurcation is bordered dorsally by a dark chromatophore band, and there is a narrow stripe of such chromatophores from the dorsal margin of the pectoral fin base across the opercle to just below the posterior margin of the eye. The red borders of the orange lateral stripe are more or less devoid of chromatophores, forming two pale stripes bordered on either side by dark chromatophores. The dorsal regions of the head and snout have a scattering of melanophores, especially dorsally. The interorbital trough has a longitudinal dark line about two chromatophores in width, which continues onto the snout as a variably wider median stripe of melanophores from the anterior pair of interorbital papillae almost to its anterior margin. Other specimens are generally similar, with considerable variation in the degree of development of the dark chromatophores on the head and body. Comparisons: Trimma erdmanni is unique among described species of the genus in its live/freshly collected colour pattern of a red to orange body with a darker lateral stripe that usually bifurcates anteriorly above the opercle, together with a horizontal pale stripe from the upper jaw to the opercle that abuts the ventral margin of the orbit. The species belongs to the grade possessing a Type B configuration of the abdominal/caudal vertebral transition. Within this grade, subsequent subdivision is usually made on the basis of presence or absence of scales in the predorsal midline. However, T. erdmanni, like a few large specimens of T. hayashii and a currently undescribed species from Cocos-Keeling, may have some scales in this region (although they are absent in most specimens, especially when < 20 mm SL). The new species may be 142

17 Richard Winterbottom distinguished from T. hayashii in having a shorter basal membrane (< 20 vs. > 50% length of fifth pelvic fin ray) and usually more pectoral fin rays (17-19, x = 18.0 vs (x = 15.8) and from the undescribed species in having 8 vs. 9 anal fin rays. The predorsal scales, if present, are confined to an area well anterior of the first dorsal fin spine and separated from it by an unscaled area, in contradistinction to the other species possessing predorsal scales, in which the posteriormost scale lies immediately anterior to the spine. Further comparisons are therefore confined to the species lacking scales in the predorsal midline. Of the remaining 28 described species without scales in the predorsal midline, the presence of 19 or more total gill rakers in T. erdmanni separates it from all but eight of them (viz. T. benjamini Winterbottom, 1996, T. bisella Winterbottom, 2000, T. cana Winterbottom, 2004, T. capostriatum (Goren, 1981), T. necopinum (Whitley, 1959) T. sostra Winterbottom, 2004, T. striatum (Herre, 1945) and T. tauro - culum Winterbottom & Zur, 2007). Trimma capo - striatum and T. striatum can be immediately separated by the presence of six thin red stripes on the head and anterior part of the body, and in possessing a full basal membrane and 2-3 branches in the fifth pelvic fin ray (vs. no thin red stripes confined to the head and anterior body, basal membrane 7-18% the length of the fifth ray, which is branched only once). Trimma benjamini is most easily separated from the new species by the thin ring of melanophores encircling the eye, with two short, ventrally directed bars onto the cheek (vs. such markings absent) and in lacking the dark lateral stripe on the body that usually bifurcates behind the eye. The two species are very similar to each other in their meristic values. Trimma bisella usually has 10 dorsal and 9 anal fin rays (vs. 9 and 8 respectively), and has an unbranched fifth pelvic fin ray, has a large white blotch on the dorsal surface of the caudal peduncle (vs. blotch absent) and four orange bars on the head (vs. bars absent) this species is apparently confined to the western Indian Ocean (Mauritius). Trimma cana and T. sostra have an unbranched fifth pelvic fin ray (vs. branched) and a translucent to white body with numerous red bars (T. cana) or blotches (T. sostra) in life (vs. such red markings absent). Trimma necopinum usually has 9 anal rays (vs. 8), a full basal membrane (vs. much reduced), and a live/fresh colour pattern of numerous red to orange spots and blotches on the head and body (vs. spots and blotches absent) this species is apparently confined to the Great Barrier Reef of Australia. Finally, T. tauroculum has an eye-diameter sized black ocellated blotch on the side of the body above the pectoral fin, and numerous smaller black spots on the nape (vs. both absent) this species has only been recorded to date from Palau, and from Ulithi Atoll in the Yap Islands (Western Carolines). Distribution: Trimma erdmanni is currently known from the Raja Ampat islands and Sulawesi in Indonesia, and the Hermit Islands and Madang, Papua New Guinea, in water depths of 0-66 m. There are also photographs of a freshly collected specimen from El Nido, Palawan Island, and of live specimens from Davao Gulf, Mindanao and Calamianes Is., Batangas in the Philippines (Fig. 6). The species has also been observed in the Lembeh 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 in Trimma (and other fishes, of course), his enthusiastic collection and documentation of specimens of this genus for the present author s research program, his friendship, and for the superb job he does for Conservation International s Indonesian Marine Program. This species has been referred to informally (in litt.) as Trimma RW sp. 68. Trimma habrum n. sp. (Figs 6, 13-15) Delicate pygmy goby Trimma RW sp. 95 Dimara et al. 2010: 621 (Raja Ampat) Material Examined: A total of 1 lot, 12 type specimens, plus two additional non-type specimens (tissue voucher specimens). The description is based on the holotype and 11 paratypes ( mm SL). Holotype: ROM 87486, 16.8 mm SL male, Indonesia, Raja Ampat, Fam Islands, Keruo Island (just east of Penemu Island), vertical wall, S; E, 70 m, clove oil, 25 January 2010, M. V. Erdmann. Paratypes: All specimens collected with the holotype: AMS I , 1, MZB 19778, 4, ROM 84881, 5, ROM 1832CS, 1, 16.6 (male). Additional (Non-type) Material. Two specimens 143 aqua vol. 17 no July 2011

18 Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature collected with ROM for genetic analysis: ROM T07710 and T07711 ( ). Diagnosis: Trimma habrum is characterized by a bony interorbital as wide as the diameter of the pupil, 8-9 scales in the predorsal midline, 14 unbranched pectoral fin rays, an unbranched fifth pelvic fin ray, no basal membrane joining the fifth pelvic fin rays across the midline, a single full row of cheek scales and sometimes one or two small scales above this row, three rows of scales covering the upper two-thirds of the opercle, and no trace of dark or black pigmentation on the caudal peduncle when preserved. The fresh colouration is also diagnostic: a pale translucent dorsum with light yellow blotches and the base of each element of the dorsal fin surrounded by a red spot, a thin red bar along the posterior margins of the hypurals, no dark pigment at all on the hypural region of the peduncle, and a darkly pigmented covering to the dorsal margins 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, second spine reaching to between base of second ray of second dorsal fin and one scale posterior to last ray when adpressed (to base of sixth ray on holotype), third to between base of spine and first ray of second dorsal fin, last ray extends posteriorly for about one-third of distance between its base and first 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 posterior element of last ray, which extends posteriorly for about one-third of distance between its base and first ventral procurrent ray; pectoral fin 14, all rays unbranched, fin reaching posteriorly to a ver- tical line above a line between urogenital papilla and anal spine; pelvic fin I 5, no frenum, basal membrane absent, first four rays with one sequential branch, fifth ray unbranched and % length of fourth (x = 46.8, SD = 2.88, n = 11), which reaches posteriorly to between bases of first to third anal-fin ray. Lateral scales 23 (n = 11), anterior transverse scales 7-8 (x = 7.2, SD = 0.25), posterior transverse scales (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 and 7-8 cycloid scales in main lower row, upper twothirds of opercle covered with three longitudinal rows of 2, 2 and 1 cycloid scales respectively (the posterodorsal scale may occasionally be weakly ctenoid); 2 vertical rows of cycloid scales on pectoral base, with 3 scales in posterior row; 4 cycloid prepelvic scales in midline; (x =11.7, SD = 0.47) circumpeduncular scales; body scales ctenoid except for cycloid scales on anterior belly midline, beneath and immediately posterior to pectoral-fin base, and first few scales in predorsal midline and around posterodorsal margin of eye (Fig. 13). Gill opening extending anteroventrally to a vertical below anterior third to middle of pupil. Upper jaw with an outer row of somewhat enlarged, spaced, slightly curved canines decreasing in size posteriorly to end of premaxilla, 2-3 irregular inner rows of small conical teeth behind symphysis decreasing to a single inner row extending to end of premaxilla; innermost row slightly larger and directed posteriorly, decreasing in size posteriorly. Lower jaw with an outer row of enlarged curved spaced canines from symphysis to bend of dentary and directed somewhat anterodorsally, 1-2 irregular Fig. 13. Dorsal view of the head of Trimma habrum (16.8 mm SL male holotype, ROM 87486). Specimen stained with cyanine blue. Photo by R. Winterbottom. aqua vol. 17 no July 2011 Fig. 14. Left lateral view of the vertebral transition of Trimma habrum, 19.2 male paratype, ROM 1832CS. Specimen stained with alizarin and alcian blue. Photo by R. Winterbottom. 144

19 Richard Winterbottom inner rows of smaller curved teeth at symphysis grading to a single row laterally and continuing to mid-height of coronoid process of dentary. Cephalic sensory papilla counts as in Table I. Tongue broadly truncate with rounded edges. Gill rakers on first arch = (x = = 16.0, SD = 0.28, 0.28, and 0.41 respectively). Anterior nasal opening a short tube extending out over upper lip, posterior nasal opening a pore with a raised rim, both protruding from slightly raised oval sac confined to anterior half of snout. Bony interorbital equal to pupil width, with a gently rounded fleshy median interorbital ridge, no dermal crest, and epaxialis musculature extending anteriorly to above a vertical with posterior margin of pupil (Fig. 13). Abdominal/caudal vertebral configuration Type A, with haemal arches of first two caudal vertebrae expanded to accommodate posterior extension of swimbladder (Fig. 14). Colour pattern (from slides of two specimens, mm SL, description based mostly on the former, Fig. 15). The 16.8 mm SL male holotype is a essentially a pale, translucent fish, with the anterior region of the dorsum suffused with light yellow, the exposed portions of the scales below the dorsal fins are thinly edged with red, there are two yellow saddles a little less than pupildiameter in width over the dorsal part of the peduncle, and one (below the second dorsal saddle) over the ventral part of the peduncle. The neural canal and upper half of the swimbladder are heavily invested with melanophores and clearly visible in lateral view; the abdomen is pale pink anteriorly grading to yellow posteriorly, the yellow intensifying into a narrow, tapering wedge posteriorly above the base of the anal fin and ending about at the anterior third of the peduncle. The whole ventral half of the body behind the abdominal cavity is reddish-pink. The snout is yellow with a reddish nasal sac, the brain case is heavily sprinkled with large dark pigment cells, and the cheek has a white stripe between the posteroventral and anteroventral margins of the eye. Below this, the cheek is yellow grading to reddish-pink posterodorsally; the opercle has a light mauve suffusion. The pectoral fin base is suffused with light purplish-red, the base of the pelvic fin is suffused with yellow. The spines of the first dorsal fin are red, each with a slightly enlarged red spot at its base, and with a yellow orange, half-pupil diameter wide yellowish orange stripe just above the base. The membranes between the first three spines are heavily sprinkled with melanophores, the fin membrane posterior to this is hyaline. The bases of all elements of the second dorsal fin are contained in red spots, the proximal half of the fin membranes is heavily suffused with yellow (more so posteriorly), and the distal half contains scattered melanophores and red and yellow chromatophores. The caudal fin contains a mix of yellow and red suffusions with scattered melanophores, and the bases of the rays and the regions adjacent to them are red, forming a thin red vertical line over the ends of the hypurals. The basal region of the anal fin has a series of red inverted triangle separated by pale areas (these do not conform to the bases of the fin elements), the fin has a basal black stripe followed by a broad yellow stripe and culminating in a distal black stripe. The pectoral fin rays are red, intensified at their bases, and the membrane is hyaline. The pelvic fin Fig. 15. Left lateral view of Trimma habrum, 16.8 mm SL male holotype, ROM 87486, Keruo Island, Raja Ampat. Photo by R. Winterbottom. 145 aqua vol. 17 no July 2011

20 Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature and fin rays are hyaline. The iris is yellow with a diffuse black margin and a horizontal, one-third pupil-diameter dark purple stripe which touches the dorsal rim of the pupil ventrally. The second photographed specimen (17.3 mm SL, ROM T7710, tissue voucher) is essentially similar, but lacks the yellow wedge from the abdomen that passes posteriorly above the base of the anal fin. Colour pattern in alcohol: off white, with darker pigmentation over the swimbladder visible through the body, as well as that covering the braincase around the posterior rim of the orbit. The dark pigmentation in the distal portions of the dorsal fins, the base and distal regions of the anal fin, and a few dark chromatophores on the dorsal surface of the snout are also visible. Comparisons: Trimma habrum belongs to the T. tevegae species complex, which is defined by the apparently apomorphic conditions of the broad bony interorbital (75% or more of pupil diameter) and a Type A abdominal/caudal vertebral transition region. Among the 10 nominal species currently assigned to this group, T. habrum differs from T. caudomaculatum Yoshino & Araga, 1975, T. griffithsi Winterbottom, 1984, T. nasa Winterbottom, 2005 and T. tevegae Cohen & Davis, 1969, as well as from T. xanthochrum n. sp. described herein, in lacking any dark pigmentation on the posterior portion of the caudal peduncle. All the remaining species except T. marinae Winterbottom, 2005, have at least some branched rays in the middle of the pectoral fin. Trimma marinae shares with T. habrum the thin vertical red bar over the ends of the hypurals in freshly collected specimens, but has an open nasal capsule lacking anterior and posterior nares (vs. a nasal sac with both nares present), fewer scales in the predorsal midline (6-8 vs. 9-10), and the bases of the elements of the dorsal fin are not surrounded by a small, dark red spot (vs. such a spot present). Distribution: Trimma habrum is currently only known from a single collection at 70 m on a vertical wall at Keruo Island (off Penemu Island), one of the Fam Islands in Raja Ampat, Indonesia (Fig. 6). Etymology: Derived from the Greek word habros, meaning delicate, dainty or graceful, in allusion to the soft and delicate shades of colour of freshly collected specimens. Trimma habrum has been referred to informally (in litt.) as T. RW sp. 95. aqua vol. 17 no July 2011 Trimma haimassum n. sp. (Figs 1 B & C, 2A, 6, 16-20) Blood-spot pygmy goby Trimma RW sp. 63 Dimara et al. 2010: 621 (Raja Ampat) 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 Raja Ampat, Indonesia) Trimma sp. 10 Kuiter & Tonozuka 2004: 706 (Indonesia) Material Examined: A total of 11 collections, 240 ( mm SL) type specimens, plus additional non-type specimens from Raja Ampat (including a tissue voucher specimen), as well as numerous specimens from several other western Pacific localities. The morphological description is based primarily on 10 males, 9 females ( mm SL, x = 27.6, SD = 1.52, ROM 85347), plus the holotype, with some details (teeth, vertebral transition) from ROM 1836CS (5, ); all specimens from the same collection. Holotype: ROM 87484, 27.3 mm SL female, E side of Barracuda Rock, 200 m N of Wayil Island ( S, E), cave with lots of sea fans & sea whips near entrance at top part, big variety of soft & hard corals just outside, m, rotenone, , field # RW10-40, 1 February 2010, R. Winterbottom, L. Katz & CI team. Paratypes: AMS I (formerly ROM 85390), 38, , SE islands off Misool, south side of Balbulol Island ( S, E), 45 m, rotenone, , field # RW10-45, 2 February 2010, M. V. Erdmann. MZB (formerly ROM 85377), 59, , SE islands off Misool, south side of Balbulol Island ( S, E), sea fans, sea whips, sponges, tunicates, hydroids, m, rotenone, , field # RW10-44, 2 February 2010, R. Winterbottom, L. Katz & CI team. ROM 84877, 5, , Keruo Island, off Penemu Island, Fam Islands ( S, E), various hard and soft corals, tunicates and sponges, m, clove oil, , field # RW10-03, 25 January 2010, R. Winterbottom & L. Katz. ROM 84884, 2, , Keruo Island off Penemu Island, Fam Islands 146

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