aqua, International Journal of

Six new of the (Percomorpha; ) 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. Email: [email protected]

Received: 30 December 2010 – Accepted: 01 April 2011 Abstract by a relatively deep body, numerous irregular scales with Recent (2010) fieldwork in the Raja Ampat Islands of about 30 lateral rows and over 15 scales in the anterior trans- Indonesia resulted in the collection of seven undescribed verse series, a fifth pelvic fin ray that branches twice dichoto- species of Trimma, six of which are described here. Two of mously, usually a somewhat elongate second dorsal spine, no these are currently only known from these islands, the others predorsal, cheek or opercular scales, a moderately wide bony are known from other parts of the western Pacific. Trimma interorbital with a fleshy median ridge between the eyes, and cheni n. sp. is distinguished from other species of Trimma in a dermal ridge anterior to the first dorsal spine. There is a red having 8-9 scales in the predorsal midline, 2-3 scales on the spot or elongate blotch above the opercle in live and fresh opercle, an elongate second dorsal spine reaching posteriorly material, and the dorsal surface of the snout has a reticulated to the bases of rays 1-4 of the second , middle pec- dark pattern, with dark transverse stripes over the dorsal mar- toral fin rays branched, fifth pelvic fin ray branched once, a gin of the orbit. The species is known from south-western dark basal stripe in the dorsal fins, scale pockets indistinctly Sulawesi north to Palawan and eastwards to the Solomon outlined with darker pigment, and, in life, two red to orange Islands. Trimma papayum n. sp. is unique among the bars across the cheek and three diffuse yellow stripes on the described species of the genus in having a one-third pupil body (most obvious along the caudal peduncle). It has been diameter black ocellated spot on and just behind the fourth recorded from , the , and Sulawesi and Flo- dorsal fin spine. It has 9 dorsal and 8 anal fin rays, a single res in Indonesia. Trimma erdmanni n. sp. has a reddish- branch point in the fifth pelvic fin ray, 10-11 anterior and 8- orange body with a dark red lateral stripe with darker borders 9 posterior transverse scale rows, 5-10 scales in the predorsal on the body which extends anteriorly onto the head, where it midline, and a single row of 3 scales on the upper margin of bifurcates, and there is a thin longitudinal light stripe below the opercle. Freshly collected specimens are orange red in the eye. It usually lacks scales in the predorsal midline, the overall colouration, with scattered diffuse yellow spots. The second spine of the first dorsal is elongated, there are 9 dorsal species is known only from Indonesia, at Maumere, Flores and 8 anal rays, a single branch in the fifth pelvic fin ray, and and Kawe Island, Raja Ampat. Trimma xanthochrum n. sp. is 19-22 gill rakers on the first gill arch. The species is known characterized by a wide interorbital region (80-100% pupil from the Raja Ampat islands, the Hermit Islands and diameter), a second dorsal spine usually reaching posteriorly Madang (), and from photographs from to between the bases of the second to third dorsal fin rays, 15- El Nido, Palawan Island and Davao Gulf, Mindanoro, 16 pectoral rays usually with 7-8 branched rays, vertical rows Philippines. Trimma habrum n. sp. has a bony interorbital of sensory papillae below eye of 2-3 papillae in rows 1-4 and width equal to the pupil diameter, 8-9 scales in the predorsal 4-5 in row 5, a caudal blotch which has a lower half about midline, 14 unbranched pectoral fin rays, an unbranched two-thirds the width of the upper half, and usually an overall fifth pelvic fin ray, no basal membrane joining the fifth pelvic yellowish body with yellow at least proximally in the caudal fin rays across the midline, usually a single full row of cheek fin. It is currently known with certainty only from the Raja scales, and scales on the upper two-thirds of the opercle. The Ampat islands. fresh colouration is diagnostic: a pale translucent dorsum with light yellow blotches and the base of each element of the Zusammenfassung dorsal fin surrounded by a red spot, a thin red bar along the Bei einer neueren Forschungsreise zum Raja-Ampat-Archi- posterior margins of the hypurals, no dark pigment at all on pel Indonesiens (2010) wurden sieben unbeschriebene the hypural region of the peduncle, and a darkly pigmented Trimma-Arten entdeckt, von denen sechs hier beschrieben covering to the dorsal margins of the swim bladder, neural werden. Zwei der neuen Arten kommen nach heutiger sheath, and the brain. It is currently known only from a sin- Kenntnis nur an diesen Inseln vor, die anderen sind auch von gle collection made at Kerou Island, Fam Islands in Raja anderen Teilen des Westpazifiks bekannt. Trimma cheni n. sp. Ampat, Indonesia. Trimma haimassum n. sp. is characterized unterscheidet sich von den anderen Trimma-Arten durch fol-

127 aqua vol. 17 no. 3 - 10 July 2011 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 Mittel- streifen über dem rückwärtigen Rand der Augenhöhle. Man linie, 2-3 Schuppen auf dem Kiemendeckel, einen verlänger- fand diese Art am südwestlichen Sulawesi nördlich von Pala- ten zweiten Rückenstachel, der nach hinten bis zur Basis der wan und nach Osten zu an den Salomoninseln. Trimma Flossenstrahlen 1-4 der zweiten Rückenflosse reicht, ver- papayum n. sp. zeichnet sich unter den beschriebenen Arten zweigte mittlere Strahlen der Brustflosse, einmal verzweigten dadurch aus, dass sich auf und direkt hinter dem vierten fünften Bauchflossenstrahl, einen dunklen Streifen am Rückenflossenstachel ein schwarzer Augenfleck mit dem Grund der Rückenflossen, undeutlich mit dunklerem Farb- Durchmesser eines Drittels der Pupille befindet. Außerdem stoff umrahmte Schuppentaschen, außerdem beim lebenden haben Vertreter dieser Art 9 Rückenflossen- und 8 Afterflos- Tier zwei rote bis orangefarbene Bänder über die Wange hin- senstrahlen, nur einen Verzweigungspunkt beim fünften weg und drei schwach gelbe Streifen auf dem Rumpf (am Bauchflossenstrahl, 10-11 vordere und 8-9 hintere quer lau- deutlichsten sichtbar auf dem Schwanzstiel). Diese Art fende Schuppenreihen, 5-10 Schuppen auf der prädorsalen konnte bei Palau, den Philippinen und Sulawesi und an der Mittellinie und eine einzelne Reihe von drei Schuppen am Insel Flores in Indonesien nachgewiesen werden. Trimma erd- oberen Rand des Kiemendeckels. Frisch gefangene Tiere sind manni n. sp. hat einen rötlich orangefarbenen Rumpf und insgesamt orangerot, mit unregelmäßig verteilten schwach einen dunkelroten Seitenstreifen mit dunkleren Rändern auf gelben Flecken. Bisher ist die Art nur von Indonesien dem Rumpf, der sich nach vorne bis zum Kopf erstreckt, wo bekannt, von der Gegend der Inseln Maumere, Flores und er sich gabelt, außerdem einen dünnen hellen Längsstreifen Kawe im Raja-Ampat-Archipel. Trimma xanthochrum n. sp. unter dem Auge. Normalerweise fehlen hier Schuppen auf ist durch folgende Merkmale gekennzeichnet: breite Interor- der prädorsalen Mittellinie, der zweite Stachel der ersten bitalregion (80-100% des Pupillendurchmessers), einen zwei- Rückenflosse ist verlängert, die Rückenflossen haben 9 Flos- ten Rückenstachel, der normalerweise nach hinten bis zwi- senstrahlen, die Afterflosse 8, der fünfte Bauchflossenstrahl schen die Ansätze des zweiten und dritten Rückenflos- verzweigt sich einmal, und der erste Kiemenbogen trägt 19- senstrahls reicht, 15-16 Brustflossenstrahlen, in der Regel mit 22 Kiemenreusen. Fundorte dieser Art sind die Inseln Raja 7-8 verzweigten Strahlen, senkrechte Reihen von Sinnespa- Ampat, die Hermit-Inseln und Madang (Papua-Neuguinea) pillen unterhalb vom Auge – mit jeweils 2-3 Papillen in den sowie, durch Fotos belegt, El Nido, Palawan-Insel und der Reihen 1 bis 4 und 4-5 Papillen in Reihe 5 –, einen Schwanz- Davao-Golf, Mindanoro, Philippinen. Bei Trimma habrum fleck, dessen untere Hälfte etwa zwei Drittel der Breite der n. sp. entspricht die Breite des Interorbitalknochens dem oberen Hälfte einnimmt, sowie normalerweise eine gelbliche Pupillendurchmesser, Vertreter dieser Art haben 8-9 Schup- Gesamtfärbung auf dem Rumpf, die sich mindestens proxi- pen auf der prädorsalen Mittellinie, 14 unverzweigte Brust- mal in einem Gelbton auf der Schwanzflosse fortsetzt. flossenstrahlen, einen unverzweigten fünften Bauchflos- Gegenwärtig kennt man diese Art mit Sicherheit nur vom senstrahl, keine Basalmembran am fünften Bauchflos- Raja-Ampat-Archipel. senstrahl über die Mittellinie hinweg, normalerweise eine ein- zelne volle Reihe von Wangenschuppen und Schuppen auf Résumé dem oberen Zweidrittel des Kiemendeckels. Außerdem ist die Une récente collecte (2010) sur les Îles Raja Ampat d’Indo - Färbung lebender oder frischtoter Exemplare artkennzeich- nésie a permis la capture de sept nouvelles espèces de Trimma, nend: der Rücken blass durchscheinend mit hellgelben dont six sont décrites ici. Deux d’entre elles sont pour l’heure Flecken, die Basis von jedem Element der Rückenflosse von connues exclusivement sur ces îles, tandis que les autres peu- einem roten Fleck umrahmt, ein dünner roter Streifen ent- vent être trouvées dans d’autres parties du Pacifique Ouest. lang den hinteren Rändern der Hypurale, überhaupt kein Trimma cheni n. sp. se distingue des autres espèces de Trimma dunkles Pigment in der hypuralen Gegend des Schwanzstiels par 8-9 écailles sur la ligne médiane prédorsale, 2-3 écailles sowie eine dunkel pigmentierte Bedeckung der dorsalen Rän- sur l’opercule, un deuxième rayon épineux étiré sur la pre- der der Schwimmblase, der Nervenscheide und des Gehirns. mière nageoire dorsale allant à l’arrière jusqu’au-dessus des Bisher ist diese neue Art nur von einem einzigen Forschungs- bases des rayons 1-4 de la seconde dorsale, des rayons de la fang an der Kerou-Insel, Fam-Inseln im Raja-Ampat-Archi- nageoire pectorale divisés au milieu, le cinquième rayon de la pel, Indonesien, bekannt. Trimma haimassum n. sp. ist pelvienne divisé une fois, une rayure sombre à la base des gekennzeichnet durch einen relativ tiefen Rumpf, zahlreiche nageoires dorsales, l’attache des écailles indistinctement mar- unregelmäßige Schuppen mit etwa 30 seitlichen Reihen und quée d’une pigmentation plus sombre, et à l’état vivant, deux über 15 Schuppen in der vorderen Querreihe, einen fünften barres rouges à orange sur la joue et trois raies jaunes diffuses Bauchflossenstrahl, der sich zweimal dichotom verzweigt, sur le corps (les plus apparentes le long du pédoncule caudal). normalerweise einen etwas verlängerten zweiten Rückensta- Elle a été recensée aux Palaos, aux Philippines, et au Sulawesi chel, das Fehlen von Schuppen prädorsal, an der Wange und et à Flores en Indonésie. Trimma erdmanni n. sp. présente un auf dem Kiemendeckel, einen mäßig breiten Interorbitalkno- corps orange rougeâtre, avec une rayure latérale rouge chen mit einem fleischigen medianen Grat zwischen den sombre, aux bordures plus foncées, qui remonte jusqu’à la Augen sowie eine Hautleiste vor dem ersten Rückenstachel. tête, où elle se divise ; là, une fine rayure longitudinale de cou- Bei lebenden oder frischtoten Exemplaren zeigt sich in roter leur claire se trouve sous l’œil. Elle ne possède habituellement Farbe ein Punkt oder länglicher Fleck oberhalb vom Kie- pas d’écailles sur la ligne médiane prédorsale, la seconde épine mendeckel, und die rückwärtige Oberfläche der Schnauze de la première dorsale est étirée, la dorsale compte 9 rayons et weist ein netzartiges dunkles Muster auf, mit dunklen Quer- l’anale 8, une seule division sur le cinquième rayon de la pel-

aqua vol. 17 no. 3 - 10 July 2011 128 Richard Winterbottom vienne et 19-22 branchiospines sur le premier arc branchial. mesure deux-tiers de la moitié supérieure, et une coloration L’espèce est connue sur les Îles Raja Ampat, les Îles Hermit et du corps globalement jaunâtre, avec au minimum du jaune à à Madang (Papouasie-Nouvelle-Guinée), et sur des photogra- proximité de la nageoire caudale. Pour l’instant, on ne l’a phies prises à El Nido (Île de Palawan) et dans le Golf de identifiée avec certitude que sur les Îles Raja Ampat. Davao (Mindanoro, Philippines). Trimma habrum n. sp. pos- sède un espacement interorbital osseux égal au diamètre de sa Sommario pupille, 8-9 écailles sur la ligne médiane prédorsale, une Recenti spedizioni nelle Isole Raja Ampat, Indonesia (2010) nageoire pectorale à 14 rayons non divisés, une nageoire pel- hanno portato al campionamento di sette specie non descritte vienne dont le cinquième rayon n’est pas divisé, une absence di Trimma, sei delle quali sono descritte in questo articolo. de membrane basale reliant le cinquième rayon de part et Due di queste sono al momento note solo in queste isole, le d’autre de la ligne médiane, habituellement une seule rangée altre anche da altre parti del Pacifico occidentale. Trimma complète de nageoires sur les joues et des écailles sur les deux cheni n. sp. si distingue dalle altre specie di Trimma per avere tiers supérieurs de l’opercule. Les tons frais de sa coloration 8-9 scaglie sulla linea mediana predorsale, 2-3 scaglie sull’op- sont diagnostiques : un dorsum pâle et translucide, avec des ercolo, la seconda spina dorsale allungata fino a raggiungere taches jaune clair et la base de chaque élément de la dorsale posteriormente la base dei raggi 1-4 della seconda pinna dor- entourée d’un point rouge, une barre rouge fine le long des sale, raggi pettorali mediani ramificati, quinto raggio pelvico bords postérieurs des hypuraux, une absence totale de pig- con una singola ramificazione, una stria basale scura sulle mentation foncée sur la région hypurale du pédoncule, et une pinne dorsali, avvallamenti delle scaglie indistintamente pigmentation sombre couvrant les bordures dorsales de la ves- delimitate da pigmento più scuro e, in vita, due barre rosso- sie natatoire, la gaine des neurones et le cerveau. Elle est pour arancio lungo le guance e sul corpo tre strie gialle diffuse (più l’instant uniquement connue sur le résultat d’une seule col- evidenti lungo il peduncolo caudale). È stata rinvenuta a lecte réalisée sur l’Île Kerou (Îles Fam, Raja Ampat, Indoné- Palau, Filippine e a Sulawesi e Flores in Indonesia. Trimma sie). Trimma haimassum n. sp. se caractérise par un corps rela- erdmanni n. sp. ha il corpo arancio-rossastro con una stria lat- tivement épais, de nombreuses écailles irrégulières avec envi- erale rosso scuro con bordi più scuri sul corpo che si esten- ron 30 rangées latérales et plus de 15 écailles dans les séries dono anteriormente sul capo, do ve si biforca, ed è presente transversales antérieures, un cinquième rayon de la pelvienne una sottile striatura longitudinale chiara sotto l’occhio. In se ramifiant deux fois de manière dichotomique, habituelle- genere non possiede scaglie predorsa li, ha la seconda spina ment une seconde épine dorsale quelque peu étirée, une della prima dorsale allungata, possiede 9 raggi dorsali e 8 absence de prédorsale, des écailles sur les joues ou les oper- anali, una singola ramificazione nel quinto raggio pelvico e cules, un espacement osseux interorbital limité avec une 19-22 rastrelli branchiali sul primo ar co branchiale. La specie excroissance charnue médiane entre les deux yeux, et une è stata raccolta alle isole Raja Ampat e a Madang e alle isole excroissance dermique avant la première épine dorsale. Un Hermit (Papua Nuova Guinea) ed è stata fotografata a El point rouge ou une tache étirée au-dessus de l’opercule est Nido, isola di Palawan e nel golfo di Davao, Mindanoro, Fil- visible sur des spécimens vivants et frais, et la surface dorsale ippine. Trimma habrum n. sp. ha una distanza interorbitale du museau présente un motif réticulé foncé, avec des rayures ossea uguale al diametro della pupilla, 8-9 scaglie predorsali, sombres transversales sur la bordure dorsale de l’orbite. L’es- 14 raggi pettorali non ramificati, quinto raggio pelvico non pèce est connue du nord du Sulawesi sud-occidental jusqu’à ramificato, membrana basale che unisce il quinto pelvico alla Palawan et l’est des Îles Salomon. Trimma papayum n. sp. est linea mediana assente, in genere una singola fila completa di unique parmi les espèces décrites dans le genre pour avoir un scaglie sulla guancia e scaglie sui due-terzi superiori dell’oper- point noir ocellé sur la pupille, d’un tiers du diamètre de celle- colo. La colorazione dell’esemplare fresco è diagnostica: il ci et un autre juste derrière le quatrième rayon épineux de la dorso è traslucente e pallido con macchie giallo chiaro e la dorsale. Elle possède une nageoire dorsale à 9 rayons et une base di ogni elemento della dorsale circondato da una mac- anale à 8, un seul point de ramification sur le cinquième chia rossa, una sottile barra rossa lungo i margini posteriori rayon de la pelvienne, 10-11 rangées d’écailles transversales degli ipurali, nessuna pigmentazione scura nella regione ipu- antérieures et 8-9 postérieures, 5-10 écailles sur la ligne rale del peduncolo e una pigmentazione scura che ricopre i médiane prédorsale, et une seule rangée de 3 écailles sur la margini dorsali della vescica natatoria, la guaina neurale e il bordure supérieure de l’opercule. Les spécimens fraîchement cervello. Al momento è nota solo da un singolo campiona- collectés ont une coloration générale rouge orange, ponctuée mento eseguito a Kerou Island, Isole Fam, Raja Ampat, de points jaunes diffus et éparpillés. L’espèce est seulement Indonesia. Trimma haimassum n. sp. si caratterizza per avere connue en Indonésie, à Maumere (Flores) et sur l’Île Kawe un corpo relativamente alto, numerose scaglie irregolari pre- (Raja Ampat). Trimma xanthochrum n. sp. est caractérisée par senti in 30 file laterali più di 15 scaglie in linea anteriore une large région interorbitale (80-100% du diamètre de la trasversale, il quinto raggio pelvico che si ramifica due volte in pupille), une deuxième épine dorsale arrivant habituellement, modo dicotomico, la seconda spina dorsale generalmente à l’arrière, entre les bases du deuxième et du troisième rayon allungata, assenza di scaglie predorsali e anche sulla guancia e de la dorsale, 15-16 rayons sur la pectorale dont habituelle- sull’opercolo, l’interorbitale moderatamente largo con una ment 7-8 sont divisés, des rangées verticales de papilles sensi- cresta mediana carnosa tra gli occhi e una cresta dermica ante- tives sous les yeux, de 2-3 papilles sur les rangées 1-4 et de 4- riore alla prima spina dorsale. È presente una macchia rossa o 5 sur la rangée 5, une tache caudale dont la moitié inférieure una chiaz za allungata sopra l’opercolo negli esemplari appena

129 aqua vol. 17 no. 3 - 10 July 2011 Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature raccolti, mentre la superficie dorsale del muso mostra un This paper describes six of those new species. Addi- motivo reticolato scuro, con striature trasversali scure sopra il tional specimens of some of these new species were margine dorsale dell’orbita. La specie è nota dal Sulawesi sud- obtained from the Raja Ampat area by Mark Erd- occidentale a nord fino a Palawan e a est fino alle Isole mann on subsequent visits. The species described Salomone. Trimma papayum n. sp. è unica tra le specie de - scritte in questo genere per avere una macchia ocellata nera here were listed under their ‘RW’ numbers by dal diametro pari a un terzo a quello della pupilla sopra e ap - Dimara et al. (2010). pena dietro la quarta spina dorsale. Possiede 9 raggi dorsali e 8 anali, un singolo punto di ramificazione nel quinto raggio METHODS pelvico, 10-11 file di scaglie trasversali anteriori e 8-9 po- Methods and the format of the descriptions fol- steriori, 5-10 scaglie predorsali e una sola fila di 3 scaglie sul low Winterbottom (2002), except that pectoral margine superiore dell’opercolo. Esemplari appena raccolti and pelvic-fin ray branching is described from pre- hanno una livrea nel complesso rosso-arancio, con mac chie served material stained with a cyanine blue solu- gialle diffusamente disseminate. La specie è nota solo in tion as outlined in Akihito et al. (1993; Akihito et Indonesia, a Maumere, Flores e Kawe Island, Raja Ampat. Trimma xanthochrum n. sp. è caratterizzata da un’ampia al. 2002: 1270). Lengths given are Standard regione interorbitale (80-100% del diametro della pupilla), Length (SL) in millimetres; SD = Standard Devia- da una seconda spina dorsale che raggiunge di solito posteri- tion; values for the holotypes are given in bold ormente lo spazio tra le basi del secondo e il terzo raggio dor- where appropriate. In the additional (non-type) sale, 15-16 raggi pettorali con generalmente 7-8 raggi ramifi- material examined sections, the catalogue number cati, file verticali di papille sensoriali sotto l’occhio formate da is followed by the number of specimens in the lot 2-3 papille nelle file 1-4 e 4-5 nella fila 5, una chiazza caudale (except where there is only a single specimen) with che ha la metà inferiore circa due terzi l’ampiezza della metà the range of SL given in parentheses. Abbreviations superiore e, infine, una colorazione generalmente giallastra con del giallo almeno prossimalmente nella pinna caudale. for repositories of material examined follow the Attualmente è nota con certezza solo alle Isole Raja Ampat. codon abbreviations published by the American Society of Ichthyologists and Herpetologists INTRODUCTION (http://199.227.217.251/files/codons.pdf). The Trimma Jordan & Seale (type species: T. caesiura letters “CS” after a ROM catalogue number denote Jordan & Seale, 1906) contains about 100 species of specimens that were cleared with trypsin and small (<30 mm SL), often colourful gobiids, primar- counter-stained with alizarin and alcian blue. The ily associated with Indo-Pacific coral reefs. Members vertebral transition type (A or B) between the of the genus may be recognized by the lack of abdominal and caudal vertebral centra is defined cephalic sensory canal pores, a much reduced and illustrated by Winterbottom (2003: Fig. 14 cephalic sensory papillae pattern, a wide gill opening (inset)), and Winterbottom & Zur (2007: Fig. 3). extending anteriorly to below the vertical limb of the Essentially, in type A, the last two abdominal ver- preopercle or, more usually, anterior to this, a lack of tebrae are of the usual perciform configuration in spicules (odontoids) on the outer gill rakers of the that they lack a bony connection across the midline first gill arch, fewer than 12 dorsal and anal fin rays, between the bases of the paired haemal arches, and and a fifth pelvic-fin ray that is equal to or more than the first (and often subsequent two to three) caudal 40% the length of the fourth pelvic-fin ray (Winter- vertebrae have a single greatly enlarged canal with bottom 1995, 2003). Although the gill rakers on the only the tips of the haemal arches fused together in first gill arch of members of this genus often have a the midline. In type B, the last two abdominal ver- row of crenulations or serrations (a series of sharply tebrae have a bridge joining the haemal arches rounded protrusions lacking any ossification) along across the midline, forming a haemal canal; the their medial margins (Fig. 1), these structures are not first caudal vertebra has broad haemal arches with considered here as the equivalent of odontoids. There a small basal haemal canal and a larger secondary are currently 64 valid species of Trimma, with canal distal to this, which forms a posteriorly taper- approximately 35 additional known undescribed ing funnel and embraces the posterior end of the species (Winterbottom & Hoese, unpublished). swimbladder. Counts and measurements were A recent (January/February, 2010) rapid survey of input directly into an Excel file with Mitutoyo dig- the Raja Ampat islands off the Bird’s Head region of ital callipers using WinWedge 3.01™ software. New Guinea sponsored by Conservation Interna- Photographs other than the portraits of fresh or tional’s Indonesia Marine Program resulted in the live specimens were produced from multiple digital collection of seven undescribed species of this genus. images taken with a Nikon D100 or D300S cam- aqua vol. 17 no. 3 - 10 July 2011 130 Richard Winterbottom

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) era attached to a Zeiss SV-8 dissecting microscope lows the orientation of, the maxilla and for a hori- at slightly incremental focal planes selected manu- zontal row across the lower cheek that is, at least in ally, or with a Canon EOS Rebel XS camera Trimma, well separated from the first row anteri- attached to a Zeiss SV-12 dissecting microscope orly. Miller (1972a) refined this nomenclature, using Zeiss AxioVision 4.8™ software and auto- using d’ for the row behind the maxilla, and reserv- matic increments. The images were then collated ing d for the row across the cheek, and this dis- into a single image using Helicon Focus 5.1™ tinction is followed in this paper. [Note: Miller (HeliconSoft). All images, drawings and image col- (1972a; 1972b) actually uses d and d 1 respectively lations were made by the author unless otherwise in the text of both papers and for the figure in the noted. Type specimens are confined to those col- 1972b paper, but employs d’ in Figure 5 of the lected in the Raja Ampat islands of Indonesia. 1972a publication. Since Sanzo (1911) consis- Cephalic sensory papillae: Nomenclature for the tently uses “prime” where appropriate for other cephalic sensory papillae follows Sanzo (1911), rows of papillae, I have opted to follow the nomen- with the exception of his row d on the lateral sur- clature used in Miller’s figure (1972a, Fig. 5)]. This face of the cheek. Sanzo (1911) uses this letter for separation of row d into two discrete entities is also both the row immediately behind, and which fol- found in both Priolepis and Egglestonichthys (Miller

131 aqua vol. 17 no. 3 - 10 July 2011 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 the anterior and posterior sections respectively). 1992). It is also present in the possibly related However, such a separation is apparently rare in Lythrypnus (Ahnelt & Bohacek (2004) - where the gobies (D. F. Hoese, in litt.). The complete nomen- authors referred to the two sections as d 1 and d 2 for clature 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 longitudi- nally 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 85033 – Fig. 2A-B. Composite image of head to show distribution image reversed); B. T. haimassum (28.8 mm SL paratype, and nomenclature of papillae. A. Trimma haimassum ROM 85347); and C. T. haimassum (27.3 SL paratype, (upper = dorsal view of left half, lower = left lateral view); ROM 1836CS). A and B stained with cyanine blue, C B. T. xanthochrum (left lateral view of cheek area). The grey cleared and stained with alizarin and alcian blue – note connecting lines in A represent the rows as counted in this lack of crenulations, indicating that they are structures paper, those in B represent parts of the nerves that stained without any calcification. in the cyanine blue preparation. aqua vol. 17 no. 3 - 10 July 2011 132 Richard Winterbottom system of nomenclature for the papillae rows, and lae. It seems probable that when homologies can be provided a table equating his names with those empirically established, a single, unified (and differ- used by previous authors (Takagi 1988: Tbl. 4). ent) nomenclature for papillae will need to be Other authors have been content to simply illus- adopted. The recent explorations of the innervations trate the papillae patterns (although Akihito et al. of the different papillae patterns in various gobioids (1988: Fig. 36) did distinguish between the trans- (e.g. Ahnelt & Bohacek 2004; Asaoka et al. 2011) is verse and longitudinal rows). Trimma has been a promising approach, but also raises some of the dif- characterized as having a reduced transverse papil- ficulties envisioned by Hoese (1983). For example, lae pattern, consisting only of longitudinal rows of the phylogenetic homology of the papillae in the papillae on the cheek (Winterbottom & Burridge rows a, c and cp may prove especially difficult to 1992; Hoese et al. 2011). One of the putative sis- resolve. Most of the head papillae have an ovoid base, ter groups of Trimma, Priolepis (Winterbottom and the long axis is nearly always oriented at right 1984; Miya, pers.comm.) has several transverse angles to the path of the nerve supplying them. rows of cheek and interorbital papillae, at least in Apparent exceptions to this in most species of the hypothesized basal members of that clade Trimma, such as rows a, c and p, where the path of (Winterbottom & Burridge 1992: Fig. 12). Most the nerve is aligned with the long axis of the bases of species of Trimma have two longitudinal rows of the papillae, may well prove to be the result of reduc- papillae (a and c) on the cheek immediately under tion to a single papilla of originally transverse rows the eye and above row d. Frequently, the anterior- that had the papillae bases transversely oriented with most papillae in rows a and c lie below the anterior respect to their immediate nerve track. border of the eye forming a vertical row of two papil- Since the papillae patterns in Trimma correspond lae, and the papillae below the posterior margin of well, for the most part, with those illustrated and the pupil form a vertical row of two or three papillae labelled by Sanzo (1911: Pl. 9, Figs 9-10) for Gobius (in the later case potentially attributable, from dorsal affinis Kolombatovic, 1891 (current status: to ventral, to one papilla from each of rows a and c Pomatoschistus pictus adriaticus Miller, 1973), Sanzo’s and a single papillae named cp by Sanzo (1911), nomenclature is employed here with the exception respectively). Ahnelt & Bohacek (2004) also employ noted above (Fig. 2A). Although the identity of all this nomenclature for the ventralmost papilla in the rows of papillae labelled in figures of Trimma in Lythrypnus, but Asaoka et al. (2011: Fig. 4) use cp for this paper appear to be positional homologues of the entire vertical row, which consists of at least five, Sanzo’s rows, they should not be construed as phylo- perhaps more, individual papillae in Pterogobius genetic homologues. Note also that the grey line elapoides. Ahnelt & Göschel (2003: Fig. 3) also use cp linking individual papillae into the named rows as for a vertical row of several papillae on the cheek in used in this paper (Fig. 2A) is not based on an exam- Quietula. One of the new species described here ination of the innervations or of the phylogenetic (Trimma xanthochrum) has two to five papillae in history of the genus, and it is probable that some each of five vertical rows in this position. Where such papillae are incorrectly attributed to or omitted from vertical rows occur, Sanzo (1911) employs numbers that row. The papillae are easily abraded from the 1 through 6 as identifiers, beginning with “1” for the surface of the head (especially those in rows b, d, z row below the anteroventral margin of the eye. The and g), and are usually difficult to see where the area fifth row, below the posterior margin of the pupil, is covered with scales. Where counts are given, they usually marks the anterior limit of row b. Given the should thus be regarded as minimum values. Those hypothesis that Trimma exhibits a reduced transverse counts exhibiting large variation probably do so pattern of cheek papillae (with T. xanthochrum pos- because some of the papillae have been lost. sessing the most plesiomorphic condition in the genus), the application of Sanzo’s (1911) nomencla- Trimma cheni n. sp. ture of rows a and c for the longitudinal papillae rows (Figs 3-6) below the eye is incorrect since these papillae repre- Face-stripe pygmy goby sent the remnants of the vertical rows 1-5 (see also Hoese et al. 2011). An alternative nomenclature is Trimma RW sp. 51 – Dimara et al. 2010: 621 given in Figure 2B, based on an interpretation that (Raja Ampat) the papillae rows below the eye in Trimma do repre- Trimma sp. 7 – Allen & Erdmann 2009: 619 sent originally more extensive vertical rows of papil- (western Bird’s Head Peninsula)

133 aqua vol. 17 no. 3 - 10 July 2011 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 2010, M. V. Erdmann. MZB 19776 (formerly specimens (10.2-22.3 mm SL), plus an additional ROM 87479), 10, 11.1-21.9, Waigeo Island, Tan- non-type specimen (tissue voucher), all from the jung Manare (00°15’26.1”S, 130°19’01.5”E), 52 m, Raja Ampat Islands, Indonesia, and other speci- clove oil, 28 May, 2010, M. V. Erdmann. ROM mens from Indonesia and the Philippines. The 85085, 4, 14.3-22.3, about 1.2 km SSE of Mutus description is based on the holotype and 19 other Island, on west side of sand spit (00°21’ 01.0”S, specimens from ROM 85161, 85138, 85179 and 130°21’25.4”E), 50 m, clove oil, 0900-0920, field 1835CS, 16.2-22.1 mm SL (x– = 19.3, SD = 1.31), # RW10-16, 28 January 2010, M. V. Erdmann. 12 males and 8 females. ROM 85138, 11, 10.2-19.2, Jef Tsiep Island, west Holotype: ROM 87481, 20.0 mm SL male, Tan- side, channel between it and small island to the jung Manare, Waigeo Island, west side at about west (00°23’05.7”S, 130°16’37.1”E), 19.8-22.9 middle of width off small cape (00°16’26.1”S, m, clove oil, 1600-1630, field # RW10-20, 28 130°19’01.5”E), 52 m, rotenone, 0750-0810, field January 2010, R. Winterbottom, L. Katz & P. # RW10-24, 29 January 2010, M. V. Erdmann. Johannes. ROM 85161, 17, 10.9-21.8, Tanjung Paratypes: AMS I.45590-001 (formerly ROM Manare, Waigeo Island, west side at about middle 85221), 3, 19.5-21.5, Wofoh Island, west coast near of width off a small cape (00°16’26.1”S, south end (00°15’21.9”S, 130°17’32.0”E), 48 m, 130°19’01.5”E), 19.8-22.9 m, rotenone, 0750- clove oil, 1610-1650, field # RW10-26, 29 January 0820, 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. 3 - 10 July 2011 134 Richard Winterbottom terbottom, R, L. Katz, P. Johannes, W. Kaka & teriorly to bases of rays 1-3-4 of second dorsal fin W. Awom. ROM 85197, 7, 13.4-19.8, collected when adpressed (x– = 2.3, SD = 1.02, n = 19), rays with the holotype. ROM 85315, 1, 17.6, S of all branched except, usually, posterior element of Misool Island, off Waaf Island (02°08’56.4”S, last ray (first ray unbranched in 2), last ray reach- 130°13’17.0”E), clove oil, 0715-0740, field # ing posteriorly 40-50% of distance from its base to RW10-36, 1 February 2010, M. V. Erdmann. first dorsal procurrent ray (x– = 48.8, SD = 3.05); ROM 1835CS (formerly ROM 85232), 2, 19.6- anal fin I 8, all but first and posterior element of 21.2, Wofoh Island, west coast near south end posteriormost ray branched; posteriormost ray 35- (00°15’21.9”S, 130°17’32.0”E), 12-16 m, clove 40-45% of distance from its base to first ventral oil, 1610-1700, field # RW10-27, 29 January procurrent ray (x– = 41.0, SD = 2.55, n = 19); pec- 2010, L. Katz. toral fin 16-17-18 (x– = 17.1, SD = 0.38), 4-5-7 Additional (Non-type) Material. Raja Ampat: (mean = 5.4, SD = 1.03) and 4-8 (x– = 5.7, SD = ROM T07737, (21.8), collected with ROM 85085 1.36) unbranched dorsal and ventral rays respec- (tissue specimen). Indonesia: Flores, ROM 62652, tively, middle rays branched, fin reaching posteri- 2 (17-19). Sulawesi, ROM 64641, 2 (19-21). orly to a vertical line between bases of anal rays 1- Palau: Ulong Pass, ROM 74805, 5 (10-18). 3 (x– = 1.3, SD = 0.56, n = 19); pelvic fin I 5, no Philippines: Cebu, ROM 49247, 8(9-19); ROM frenum, basal membrane reduced and 10-15 % 53114, 3 (20-21). Negros Oriental, ROM 53112, length of fifth ray (x– = 13.0, SD = 1.91), first four 9 (12-22); ROM 53113, (18). Siquijor, ROM rays with one sequential branch, fifth ray branched 53107, (19); ROM 53108, 6 (19-22); ROM once and 48-52-56% length of fourth (x– = 52.6, 53109, 8 (16-21); ROM 53110, 11 (10-22); ROM SD = 2.50, n = 18), fourth ray reaching posteriorly 53111, 8 (21-24). to between bases of anal rays 1-3.5-6 (x– = 4.0, SD Diagnosis: Trimma cheni has 8-9 scales in the = 1.24, n = 17). Lateral scales 23; anterior trans- predorsal midline, a row of 2-3 usually ctenoid verse scales 7-8-8.5 (x– = 8.1, SD = 0.37); posterior scales along the upper border of the opercle, no transverse scales 7-8-8.5 (x– = 7.5, SD = 0.41); pre- cheek scales, a dark basal stripe in the dorsal fins, dorsal scales 8-9 (x– = 8.2, SD = 0.39); no scales on an elongate second dorsal spine reaching posteri- cheek; opercle with single row of 2-3 usually orly to the bases of rays 1-4 of the second dorsal ctenoid scales; pectoral base with usually 3 vertical fin, the middle rays of the pectoral fin branched, a rows of scales and 4 cycloid scales in posterior row; fifth pelvic fin ray which branches once dichoto- 5-7-8 (x– = 6.4, SD = 0.79) cycloid prepelvic scales mously, the basal membrane connecting the inner (in midline anterior to basal membrane); 11-12 (x– margins of the fifth pelvic fins rays is less than 15% = 11.9, SD = 0.3) circumpeduncular scales; body the length of the fifth ray, the scale pockets are not scales ctenoid except for cycloid scales on anterior distinctly outlined with darker pigment, and, in belly midline and on, beneath and just posterior to life, there are two red to orange bars across the pectoral fin base; body scales extending anteriorly cheek and three diffuse yellow stripes on the body to just behind eye. Gill opening extending (most obvious along the caudal peduncle). anteroventrally to a vertical below anterior one- Description: Dorsal fins VI + I 8-9 (x– = 9.0, SD third of pupil. Upper jaw with outer row of curved, = 0.22), second third spine longest, reaching pos- 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. 3 - 10 July 2011 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 Bony interorbital 31-35-44% (x– = 38.1, SD = 4.08) not much larger than inner rows; two to three irreg- pupil width, with broad moderately developed U- ular inner rows of smaller teeth almost reaching dis- shaped interorbital furrow and shallow postorbital tal tip of premaxilla, innermost row of slender, groove (or none). No ridge of tissue (or dermal slightly curved, spaced teeth almost as long as outer crest) extending anterior to first dorsal spine. Epax- row, from symphysis to bend of jaw in males and ialis musculature extending anteriorly to above ver- some females. Lower jaw with outer row of 4-5 tical with posterior margin of pupil. Last two curved, enlarged, spaced canines reaching to bend abdominal vertebrae with bridge of bone connect- of dentary, several irregular inner rows of slightly ing haemal arches just distal to their bases, forming curved caniform teeth half size of outer, becoming ‘haemal canal’ bounded dorsally by centrum. First reduced in size and less curved posteriorly and caudal vertebra with two canals, a small proximal tapering to form single row of straight conical teeth one at base of haemal arches, which fuse in midline at beginning of coronoid process; innermost row of briefly before diverging again to form posteriorly enlarged slightly curved teeth at symphysis, increas- directed funnel-like second arch. Haemal arches ing in size along jaw (equal in size to outermost row then fusing again in midline to form haemal spine just behind bend of dentary, then gradually decreas- (vertebral transition Type B). ing in size to dorsal tip of coronoid process (becom- Colour pattern (from images of four freshly ing equal in size to teeth in inner rows). Cephalic collected specimens, 17.6-22.3 mm SL). A 22.3 sensory papillae row counts given in Table I. mm SL male (Fig. 3) has a mostly red head, darker Tongue truncate with rounded edges. Gill rakers on on the snout, with two lighter and more intense first arch 3-4 + 15-16 = 18-19-20 (x– = 19.2, SD = red bars with irregular diffuse yellowish margins on 0.59). Anterior nasal opening a short tube extend- the cheek below the anterior and middle of the eye, ing out over upper lip, posterior nasal opening a and another yellow-brown bar over the vertical pore with raised rim, both protruding from slightly limb of the preopercle, the dorsal half of the oper- raised oval sac confined to anterior half of snout. cle 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. erd- manni; ✪ T. haimassum; ♦ T. papayum; ★ T. xanthochrum. aqua vol. 17 no. 3 - 10 July 2011 136 Richard Winterbottom of the branchiostegal membrane immediately adja- ent. The dark basal stripes in the dorsal and anal cent to the opercle and subopercle is orange. The fins remain obvious, the dark stripe at the mid- iris is red grading to yellow orange ventrally, with region of the second dorsal fin is relatively well an anteroventral to posterodorsal dark purple stripe defined, and three diffuse dark stripes run parallel across the pupil; the dorsal surface of the orbit has to the rays of the caudal fin. The dorsal rim of the several alternating red and white stripes passing pectoral fin base is often somewhat darker than the across the midline. The body is yellowish, grading surroundings, appearing as a diffuse dark streak. to pink on the peduncle, with three vague yellow Comparisons: Trimma cheni belongs to a group of stripes (most obvious on the peduncle), with heav- species which has a scaled predorsal midline, no ily scattered dark chromatophores, especially on cheek scales, a scaled opercle, at least the middle the dorsum and along the scale pockets and scale rays of the pectoral fin are branched, and a dark margins. The membranes of the dorsal fins are basal stripe in the dorsal fins is present (Winterbot- densely speckled with melanophores, except for a tom 2006). Within this group, T. cheni appears half-pupil diameter wide yellow stripe just above closest morphologically to T. milta Winterbottom, the base and, above this, a second but more diffuse 2002, in having a poorly developed postorbital yellow stripe. The membranes of the caudal fin are trough or none and in possessing a single row of 2- similarly invested with melanophores, with several 3 scales on the opercle. However, T. milta does not streaks and blotches of yellow separated by three have the second dorsal spine elongated, has an darker diffuse stripes. The distal margin of the anal unbranched fifth pelvic fin ray, has the scale pock- fin is dusky, and there is a broad (about one-and-a- ets on the body lightly outlined with darker pig- half pupil diameters wide) yellow stripe with dif- ment, and lacks the conspicuous red to orange bars fuse margins and a dark basal stripe. The pectoral on the cheek as well as the three yellow stripes on and pelvic fin rays are yellow or orange, with hya- the body. Trimma anthrenum Winterbottom, 2006, line membranes, the bases of the fins are light tan differs in lacking opercular scales and elongate sec- with many scattered dark brown chromatophores. ond dorsal spine, in possessing a well developed A 20.1 mm SL male is similar, but with a darker postorbital trough, and in live colouration (plain and greyer body, while a 20.0 mm SL male is paler, yellow with a distinctively coloured iris (see Win- with a generally more yellow cast. A specimen from terbottom 2006: Figs 1-2). Trimma preclarum Win- El Nido, Philippines photographed underwater terbottom 2006 differs from T. cheni in lacking (Fig. 4) has a greyish background with diffuse yel- scales on the opercle, in possessing a well developed low-green markings on the body, three bars on the postorbital trough, and in lacking the two red to lower part of the head (two below the eye, the third orange bars on the cheek, although this species does over the vertical limb of the preopercle), which have three yellow stripes on the posterior part of the grade from orange yellow dorsally into the green- body. These stripes are also evident in T. hayashii ish-yellow ventrally in the region of the lower jaw Hagiwara & Winterbottom, 2007, which lacks and branchiostegals. There are two short oblique scales in the predorsal midline (or, if scales are pre- orange bars passing posterodorsally from the poste- sent, they are represented one or two scales crossing rior margin of the upper orbit, and three short the midline between the eye and the first dorsal fin orange to red transverse bars across the dorsal mar- origin) and on the opercle, and has a distinctive gin of the eye. Other specimens photographed black ocellated spot on the branchiostegal mem- underwater have a redder head and either a green- branes below the vertical limb of the preopercle. ish-yellow or brownish body. A 17.6 mm SL juve- Distribution: Trimma cheni is currently known nile (Fig. 5) has a relatively dark body, two dark red from the Raja Ampat islands, and from Palau, the bars on a grey-pink cheek, and the stripes in the Philippines and south-west to Sulawesi and Flores fins are reddish rather than yellow. in Indonesia (Fig. 6) at depths of 5-52 m. Colour pattern in alcohol: straw- Etymology: The species is named for I.-S. Chen, coloured with a heavy scattering of brown chro- National Taiwan Ocean University, Keelung, Tai- matophores and melanophores on the body, nape wan, in acknowledgment of his numerous publica- and pectoral fin base. The opercle has numerous tions on Indo-Pacific reef fishes in general, and densely scattered melanophores. The cheek is paler, gobies in particular. This species has been referred and two diffuse lighter bars (corresponding to the to as Trimma RW sp. 51. red bars in fresh material) may or may not be appar-

137 aqua vol. 17 no. 3 - 10 July 2011 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. Kawe Island, SW bay (00°05’15.2”S, 130°07’ (Figs. 1A, 6-12) 25.3”E), 50 m, clove oil. M. V. Erdmann, 26 Jan- Erdmann’s pygmy goby uary 2010. Paratypes: AMS I.45591-001 (formerly ROM Trimma anaima Winterbottom 2000 – Allen & 85330), 1, 23.1, Kepotsol Island, east side Erdmann 2009: 619 (in part, Bird’s Head Penin- (02°09’32.1” S, 130°17’34.0” E), 66 m, clove oil, sula) field # RW10-38, 0830-0850, 1 February 2010, Trimma RW sp. 68 – Dimara et al. 2010: 621 M. V. Erdmann. MZB 19777 (formerly ROM (Raja Ampat) 87460), 7, 11.2-22.5, Waigeo Island, Tanjung Trimma sp. – Allen et al. 2003: 329 (lower right), Manare (00°15’26.1”S, 130°19’01.5”E), 52 m, Philippines clove oil, no field No., 28 May, 2010, M. V. Erd- mann. ROM 85033, 22, 10.2-26.2, collected with Material Examined: A total of 7 lots, 64 type the holotype. ROM 85144, 8, 12.3-23.5, Jef Tsiep specimens, plus four additional non-type speci- Island, west side, channel between it and small mens (tissue vouchers), and three additional lots of island to the west (00°23’05.7”S, 130°16’37.1”E), non-type specimens from Papua New Guinea. All 42 m, clove oil, field # RW10-21, 1600-1620, 28 type specimens collected at the Raja Ampat January 2010, M. V. Erdmann. ROM 85186, 16, Islands, Indonesia. Description based on up to 24 10.5-22.3, Tanjung Manare, Waigeo Island, west specimens from ROM 85033, 85144, 85186, side at about middle of width off small cape, 87460, 87484 and 1837CS (18.2-26.2 mm SL, 12 sponges, hard corals (00°16’26.1”S, 130°19’01.5” males, 12 females). E), 52 m, 1 kg powdered rotenone & detergent, Holotype: ROM 87482, 23.5 mm SL male, field # RW10-24, 0750-0810, 29 January 2010, M. V. Erdmann. ROM 85393, 7, 15.2-22.2, SE islands 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, west coast 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 and T07744, 2 (7.0-21.1), collected with ROM 85144. Papua New Guinea. Hermit Islands: USNM 243952, 17 (13-21. Madang: WAM P.30358-008, (19.0); WAM P. 30369-011, (21.2). Diagnosis: Trimma erdmanni differs most tren- chantly 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 Fig. 7A-B. Head of Trimma erdmanni (19.2 mm SL female dorsal fin by an unscaled area. The second (and paratype, ROM 85033) in: A. left lateral and B. dorsal third to some degree) spines of the first dorsal are view. Specimen stained with cyanine blue. Photo by R. elongated, there are 9 dorsal and 8 anal rays, the Winterbottom. middle rays of the pectoral fin are branched, there aqua vol. 17 no. 3 - 10 July 2011 138 Richard Winterbottom is a single branch in the fifth pelvic fin ray, the of distance between its base and first dorsal procur- basal membrane is 7-18% of the length of the fifth rent caudal fin ray; anal fin I 8 (n = 24), all but first fin ray and there are 19-22 gill rakers on the outer and posterior element of posteriormost ray margin of the first gill arch. branched, last ray extending between 40-80% (x– = Description: Dorsal fins VI + I 9, second and 60.7, SD = 12.4) of distance between its base and third spines longest, second spine reaching to first ventral procurrent caudal fin ray; pectoral fin between bases of second to eighth rays of second 17-18-19 (x– = 18.0, SD = 0.61), 3-4 (x– = 3.4, SD dorsal fin, third spine reaching to between bases of = 0.49) and 2-4-7 (x– = 3.4, SD = 1.00) unbranched dorsal spine and fourth ray of second dorsal fin dorsal and ventral rays respectively, middle rays when adpressed, first ray of second dorsal fin usu- branched, fin reaching posteriorly to a vertical line ally unbranched (branched in 5, SD = 0.41), pos- between anterior margin of urogenital papilla and terior element of last ray unbranched, last ray base of first ray of anal fin; pelvic fin I 5, no extending between 50-100% (x– = 74.2, SD = 16.0) frenum, basal membrane poorly developed, 7-17-

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. 3 - 10 July 2011 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 tial papilla and base of first anal-fin ray. Lateral one or two sequential branch points, fifth ray scales 23-24 (x– = 23.1, SD = 0.30, n = 20), ante- branched once and 52-61-70% length of fourth (x– rior transverse scales 8-11 (x– = 9.3, SD = 0.73, n = = 64.5, SD = 4.44, n = 20), fourth ray reaching 19), posterior transverse scales 8-9, (x– = 8.1, SD = posteriorly to between anterior margin of urogen- 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. Winter- bottom.

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. 3 - 10 July 2011 140 Richard Winterbottom in two specimens with one or two cycloid scales trench. No dermal crest anterior to first dorsal fin. respectively on upper part of opercle); midline of Epaxialis musculature extending anteriorly to predorsal usually scaleless, but with 6 scales in about a vertical with posterior margin of pupil holotype, 4 in a 20.9 mm SL female, and 6 scales (Fig. 7B). Abdominal/caudal vertebral pattern is in front of first dorsal spine followed anteriorly by Type B, but haemal arch of second caudal vertebra a naked area and then 2 more scales across midline is somewhat enlarged. in a 26.2 mm SL male; usually 4 cycloid scales in Colour pattern (from slides of six freshly posterior vertical row on pectoral base (once 5), collected specimens, 10.2-26.2 mm SL). A 26.2 with two vertical rows of scales anterior to this; 5- mm SL male (ROM 85033, Fig. 8) has an overall 6-8 (x– = 7.2, SD = 0.73, n = 20) prepelvic cycloid pinkish hue, darker on the dorsum, dominated by scales in midline anterior to pelvic-fin base; 12 cir- a slightly more than pupil-width midlateral orange cumpeduncular scales; 8-9 (x– = 8.1, SD = 0.31, n stripe bordered with red, which continues anteri- = 18) in midline between base of last anal ray and orly onto the head behind the eye in a shallow “>”- first procurrent caudal ray; body scales ctenoid shape, the lower limb touching the ventral margin except for cycloid scales on anterior belly midline, of the eye, the upper limb to a point in line with beneath and immediately posterior to pectoral-fin the dorsal margin of the pupil. In this specimen, base, anterior scales on sides of nape, and along the two arms of the “>” are joined across the inter- base of first dorsal fin; generally, body scales extend space by a narrow red-orange bridge, the interspace anteriorly on sides of nape to between a vertical is pinkish-grey. The red-orange colouration contin- above anterior margin of opercle and just posterior ues onto the snout anterior to the orbit and the to posterior margin of eye (Fig. 7). Gill opening anterior part of both upper and lower lips. The extending anteroventrally to a vertical below pupil. cheek is pale pinkish grey, bordered dorsally by a Upper jaw with outer row of curved, spaced, thin (about one-fifth pupil-diameter in width) enlarged canines, decreasing in size posteriorly to bluish-white stripe which ends at the vertical limb distal tip of premaxilla, a band of several irregular of the preopercle. The pectoral fin base and isth- rows of small conical teeth at symphysis, number mus are pale grey. A reddish-orange stripe curves of rows decreasing posterolaterally and ending just posterodorsally from the upper margin of the orbit, posterior to bend of dentary, innermost row ending above the middle of the opercle. The nape slightly larger and directed posteriorly at symph- is darker red than most of the rest of the body, and ysis, decreasing in size and becoming vertically ori- the dorsal half of the caudal peduncle is somewhat ented, and extending to distal tip of premaxilla. less so, with the space beneath the dorsal fins pale Lower jaw with an outer row of curved, spaced, pink. Most of the scales on the dorsal half of the enlarged canines ending at bend of dentary, several body have a very thin margin of red chro- rows of small, slightly curved conical teeth at sym- matophores along their posterior margins. The physis, grading to a single row at bend of dentary rows above the lateral scales and the row beneath it and continuing to lower coronoid process as a sin- have a wider band of darker red pigmentation. The gle outer row, innermost row at bend of dentary abdomen is pale pink, which grades into pale red abruptly larger than other inner teeth (twice the posteriorly. The midlateral stripe is flanked by dif- height) in males, decreasing somewhat in size pos- fuse darker stripes about equal in width, formed by teriorly and ending at the upper margin of the scattered dark chromatophores, which also under- coronoid process. Cephalic sensory papillae as in lie the red stripe, especially in the abdominal Table I and Fig. 7. Tongue truncate with rounded region. The elements of the first dorsal fin each edges. Gill rakers on first arch 4-5 + 14-16-17 = have a yellow spot just distal to the base, and there 19-21-22 (x– = 20.4, SD = 0.73, n = 21), gill rakers is a very diffuse, half-pupil diameter dark stripe with obvious fleshy crenulations (Fig. 1A). Ante- above the spots, with a scattering of melanophores rior nasal opening a short tube extending out over in the fin membrane above this. The anterior ele- upper lip, posterior nasal opening a pore with a ments of the second dorsal fin have similar yellow raised rim, both protruding from slightly raised spots, and melanophores are densely scattered oval sac confined to anterior half of snout. Bony throughout the fin membrane. The central rays of interorbital 32-50-52% pupil width (x– = 42.6, SD the caudal fin have splotches of yellow a little dis- = 5.26, n = 20), with broadly U-shaped interorbital tal to their bases, with an overall rosy hue (espe- trench with sloping sides and no postorbital cially proximally and centrally), the membranes of

141 aqua vol. 17 no. 3 - 10 July 2011 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 wedge over the hypural region and anterior bases of numerous scattered melanophores. The anal fin the caudal fin rays. has a faint pinkish hue proximally with many Colour pattern in alcohol. The 26.2 melanophores scattered in the membranes distally. mm SL male is straw coloured, including all areas Pectoral and pelvic fin membranes are hyaline, the that are red, orange or yellow in fresh specimens; rays of the median fins and the pectoral fin are pale however, most of the dark chromatophores and the red. The iris is golden-yellow, with scattered red melanophores remain obvious. The centre of the pigmentation, and heavily invested with dark pig- body stripe above the pectoral fin has a diffuse pos- mentation (especially dorsally), with a thin white teriorly tapering darker stripe formed by scattered inner rim, and a dark blue band which crosses the small chromatophores, and the margins of the iris from the anteroventral to the posterodorsal main stripe are bordered dorsally and ventrally by margins. A live specimen (Fig. 9) is generally simi- a dark diffuse pupil-width stripe of melanophores lar, although the interspace of the “>” part of the that tapers somewhat posteriorly and ends at the lateral stripe on the head almost totally occluded, hypural plate. A few small melanophores are scat- and the yellow spots in the dorsal fins are hardly tered over the body. The V-shaped anterior bifur- discernable. The snout is orange, with the posterior cation of the stripe is not apparent anteriorly, but nasal opening in a small white spot, a similar spot each arm of the bifurcation is bordered dorsally by just in fron of the eye, a median white stripe from a dark chromatophore band, and there is a narrow the anterior interorbital region to the level of the stripe of such chromatophores from the dorsal posterior nares and another similar short stripe margin of the pectoral fin base across the opercle to from the posterior interorbital region over the dor- just below the posterior margin of the eye. The red sum. In a 20.3 mm SL female (from the same lot), borders of the orange lateral stripe are more or less the mid-part of the lateral stripe is more yellow devoid of chromatophores, forming two pale than orange, and the yellow spots of the dorsal fins stripes bordered on either side by dark chro- form stripes. A 23.5mm male (Fig. 10, holotype) matophores. The dorsal regions of the head and has the whole body more heavily invested with snout have a scattering of melanophores, especially melanophores, especially anteriorly, and the pinks dorsally. The interorbital trough has a longitudinal and oranges are replaced by yellow. The interspace dark line about two chromatophores in width, of the “>” on the side of the head is almost which continues onto the snout as a variably wider occluded by the two arms of the stripe, the yellow median stripe of melanophores from the anterior spots in the first dorsal fin and anterior part of the pair of interorbital papillae almost to its anterior second dorsal fin form diffuse stripes, the dark margin. Other specimens are generally similar, stripe above this is less discernable, but there is a with considerable variation in the degree of devel- diffuse dark stripe above the yellow in the second opment of the dark chromatophores on the head dorsal fin; in both fins, there is a much greater con- and body. centration of melanophores. The basal region of Comparisons: Trimma erdmanni is unique the anal fin is pale yellow rather than pink. A 14.3 among described species of the genus in its mm SL juvenile is very dark overall, with densely live/freshly collected colour pattern of a red to scattered dark chromatophores on the head and orange body with a darker lateral stripe that usually body (Fig. 11). The orange-red lateral stripe is only bifurcates anteriorly above the opercle, together discernable on the head and scarcely at all on the with a horizontal pale stripe from the upper jaw to body, and the yellow stripes in the dorsal fins are the opercle that abuts the ventral margin of the distinct. A 10.2 mm SL juvenile (Fig. 12) has a orbit. The species belongs to the grade possessing a translucent body and head except for the dark Type B configuration of the abdominal/caudal ver- chromatophores, which are accentuated along the tebral transition. Within this grade, subsequent midlateral orange-yellow stripe. The upper lip, subdivision is usually made on the basis of presence snout, top of the head and “>” arms of the lateral or absence of scales in the predorsal midline. How- stripe are yellow, and the median fins have a scat- ever, T. erdmanni, like a few large specimens of T. tering of melanophores. A live specimen pho- hayashii and a currently undescribed species from tographed in Davao Gulf, Mindanao by Arthur Cocos-Keeling, may have some scales in this region Bos has the posterior region of the lateral red stripe (although they are absent in most specimens, espe- expanded posteroventrally to a blunt triangular cially when < 20 mm SL). The new species may be aqua vol. 17 no. 3 - 10 July 2011 142 Richard Winterbottom distinguished from T. hayashii in having a shorter and blotches absent) – this species is apparently basal membrane (< 20 vs. > 50% length of fifth confined to the Great Barrier Reef of . pelvic fin ray) and usually more pectoral fin rays Finally, T. tauroculum has an eye-diameter sized (17-19, –x = 18.0 vs. 14-17 (x– = 15.8) and from the black ocellated blotch on the side of the body undescribed species in having 8 vs. 9 anal fin rays. above the pectoral fin, and numerous smaller black The predorsal scales, if present, are confined to an spots on the nape (vs. both absent) – this species area well anterior of the first dorsal fin spine and has only been recorded to date from Palau, and separated from it by an unscaled area, in contradis- from Ulithi Atoll in the Yap Islands (Western Car- tinction to the other species possessing predorsal olines). scales, in which the posteriormost scale lies imme- Distribution: Trimma erdmanni is currently diately anterior to the spine. Further comparisons known from the Raja Ampat islands and Sulawesi are therefore confined to the species lacking scales in Indonesia, and the Hermit Islands and Madang, in the predorsal midline. Of the remaining 28 Papua New Guinea, in water depths of 0-66 m. described species without scales in the predorsal There are also photographs of a freshly collected midline, the presence of 19 or more total gill rak- specimen from El Nido, Palawan Island, and of live ers in T. erdmanni separates it from all but eight of specimens from Davao Gulf, Mindanao and them (viz. T. benjamini Winterbottom, 1996, T. Calamianes Is., Batangas in the Philippines (Fig. bisella Winterbottom, 2000, T. cana Winterbot- 6). The species has also been observed in the Lem- tom, 2004, T. capostriatum (Goren, 1981), T. beh Strait area of Sulawesi (G. R. Allen, pers. necopinum (Whitley, 1959) T. sostra Winterbot- comm., shown with a “?” mark in Fig. 6). tom, 2004, T. striatum (Herre, 1945) and T. tauro - Etymology: The species is named for Mark V. culum Winterbottom & Zur, 2007). Trimma capo- Erdmann in appreciation of his deep interest in striatum and T. striatum can be immediately sepa- Trimma (and other fishes, of course), his enthusi- rated by the presence of six thin red stripes on the astic collection and documentation of specimens of head and anterior part of the body, and in possess- this genus for the present author’s research pro- ing a full basal membrane and 2-3 branches in the gram, his friendship, and for the superb job he fifth pelvic fin ray (vs. no thin red stripes confined does for Conservation International’s Indonesian to the head and anterior body, basal membrane 7- Marine Program. This species has been referred to 18% the length of the fifth ray, which is branched informally (in litt.) as Trimma RW sp. 68. only once). Trimma benjamini is most easily sepa- rated from the new species by the thin ring of Trimma habrum n. sp. melanophores encircling the eye, with two short, (Figs 6, 13-15) ventrally directed bars onto the cheek (vs. such Delicate pygmy goby markings absent) and in lacking the dark lateral stripe on the body that usually bifurcates behind Trimma RW sp. 95 – Dimara et al. 2010: 621 the eye. The two species are very similar to each (Raja Ampat) other in their meristic values. Trimma bisella usu- ally has 10 dorsal and 9 anal fin rays (vs. 9 and 8 Material Examined: A total of 1 lot, 12 type spec- respectively), and has an unbranched fifth pelvic imens, plus two additional non-type specimens fin ray, has a large white blotch on the dorsal sur- (tissue voucher specimens). The description is face of the caudal peduncle (vs. blotch absent) and based on the holotype and 11 paratypes (15.5-17.7 four orange bars on the head (vs. bars absent) – this mm SL). species is apparently confined to the western Holotype: ROM 87486, 16.8 mm SL male, Indian Ocean (Mauritius). Trimma cana and T. sos- Indonesia, Raja Ampat, Fam Islands, Keruo Island tra have an unbranched fifth pelvic fin ray (vs. (just east of Penemu Island), vertical wall, branched) and a translucent to white body with 0°35’15.4”S; 130°17’41.1”E, 70 m, clove oil, 25 numerous red bars (T. cana) or blotches (T. sostra) January 2010, M. V. Erdmann. in life (vs. such red markings absent). Trimma Paratypes: All specimens collected with the holo- necopinum usually has 9 anal rays (vs. 8), a full type: AMS I.45592-00, 1, 17.5. MZB 19778, 4, basal membrane (vs. much reduced), and a 15.5-17.6. ROM 84881, 5, 16.3-17.7. ROM live/fresh colour pattern of numerous red to orange 1832CS, 1, 16.6 (male). spots and blotches on the head and body (vs. spots Additional (Non-type) Material. Two specimens

143 aqua vol. 17 no. 3 - 10 July 2011 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 84881 for genetic analysis: tical line above a line between urogenital papilla ROM T07710 and T07711 (17.3-18.2). and anal spine; pelvic fin I 5, no frenum, basal Diagnosis: Trimma habrum is characterized by a membrane absent, first four rays with one sequen- bony interorbital as wide as the diameter of the tial branch, fifth ray unbranched and 44-46-52% pupil, 8-9 scales in the predorsal midline, 14 length of fourth (x– = 46.8, SD = 2.88, n = 11), unbranched pectoral fin rays, an unbranched fifth which reaches posteriorly to between bases of first pelvic fin ray, no basal membrane joining the fifth to third anal-fin ray. Lateral scales 23 (n = 11), pelvic fin rays across the midline, a single full row anterior transverse scales 7-8 (x– = 7.2, SD = 0.25), of cheek scales and sometimes one or two small posterior transverse scales 7-7.5 (x– = 7.0, SD = scales above this row, three rows of scales covering 0.14), scales across predorsal midline 9-10 (x– = 9.2, the upper two-thirds of the opercle, and no trace of SD = 0.37), cheek with 1-2 scales in upper row and dark or black pigmentation on the caudal peduncle 7-8 cycloid scales in main lower row, upper two- when preserved. The fresh colouration is also diag- thirds of opercle covered with three longitudinal nostic: a pale translucent dorsum with light yellow rows of 2, 2 and 1 cycloid scales respectively (the blotches and the base of each element of the dorsal posterodorsal scale may occasionally be weakly fin surrounded by a red spot, a thin red bar along ctenoid); 2 vertical rows of cycloid scales on pec- the posterior margins of the hypurals, no dark pig- toral base, with 3 scales in posterior row; 4 cycloid ment at all on the hypural region of the peduncle, prepelvic scales in midline; 11-12 (x– =11.7, SD = and a darkly pigmented covering to the dorsal mar- 0.47) circumpeduncular scales; body scales ctenoid gins of the abdominal cavity and the brain. except for cycloid scales on anterior belly midline, Description: Dorsal fins VI + I 8-9 (x– = 8.8, SD beneath and immediately posterior to pectoral-fin = 0.37), second and third spines longest, second base, and first few scales in predorsal midline and spine reaching to between base of second ray of around posterodorsal margin of eye (Fig. 13). Gill second dorsal fin and one scale posterior to last ray opening extending anteroventrally to a vertical when adpressed (to base of sixth ray on holotype), below anterior third to middle of pupil. Upper jaw third to between base of spine and first ray of sec- with an outer row of somewhat enlarged, spaced, ond dorsal fin, last ray extends posteriorly for slightly curved canines decreasing in size posteri- about one-third of distance between its base and orly to end of premaxilla, 2-3 irregular inner rows first dorsal procurrent ray, all rays branched except, of small conical teeth behind symphysis decreasing usually, posterior element of last ray; anal fin I 8-9 to a single inner row extending to end of premax- (x– = 8.1, SD = 0.28), all rays branched except pos- illa; innermost row slightly larger and directed pos- terior element of last ray, which extends posteriorly teriorly, decreasing in size posteriorly. Lower jaw for about one-third of distance between its base with an outer row of enlarged curved spaced and first ventral procurrent ray; pectoral fin 14, all canines from symphysis to bend of dentary and rays unbranched, fin reaching posteriorly to a ver- directed somewhat anterodorsally, 1-2 irregular

Fig. 14. Left lateral view of the vertebral transition of Fig. 13. Dorsal view of the head of Trimma habrum (16.8 Trimma habrum, 19.2 male paratype, ROM 1832CS. mm SL male holotype, ROM 87486). Specimen stained Specimen stained with alizarin and alcian blue. Photo by with cyanine blue. Photo by R. Winterbottom. R. Winterbottom. aqua vol. 17 no. 3 - 10 July 2011 144 Richard Winterbottom inner rows of smaller curved teeth at symphysis the base of the anal fin and ending about at the grading to a single row laterally and continuing to anterior third of the peduncle. The whole ventral mid-height of coronoid process of dentary. half of the body behind the abdominal cavity is Cephalic sensory papilla counts as in Table I. reddish-pink. The snout is yellow with a reddish Tongue broadly truncate with rounded edges. Gill nasal sac, the brain case is heavily sprinkled with rakers on first arch 3-4 + 12-13 = 15-16-17 (x– = large dark pigment cells, and the cheek has a white 3.1 + 12.9 = 16.0, SD = 0.28, 0.28, and 0.41 stripe between the posteroventral and anteroventral respectively). Anterior nasal opening a short tube margins of the eye. Below this, the cheek is yellow extending out over upper lip, posterior nasal open- grading to reddish-pink posterodorsally; the oper- ing a pore with a raised rim, both protruding from cle has a light mauve suffusion. The pectoral fin slightly raised oval sac confined to anterior half of base is suffused with light purplish-red, the base of snout. Bony interorbital equal to pupil width, with the pelvic fin is suffused with yellow. The spines of a gently rounded fleshy median interorbital ridge, the first dorsal fin are red, each with a slightly no dermal crest, and epaxialis musculature extend- enlarged red spot at its base, and with a yellow ing anteriorly to above a vertical with posterior orange, half-pupil diameter wide yellowish orange margin of pupil (Fig. 13). Abdominal/caudal verte- stripe just above the base. The membranes between bral configuration Type A, with haemal arches of the first three spines are heavily sprinkled with first two caudal vertebrae expanded to accommo- melanophores, the fin membrane posterior to this date posterior extension of swimbladder (Fig. 14). is hyaline. The bases of all elements of the second Colour pattern (from slides of two speci- dorsal fin are contained in red spots, the proximal mens, 16.8-17.3 mm SL, description based mostly half of the fin membranes is heavily suffused with on the former, Fig. 15). The 16.8 mm SL male yellow (more so posteriorly), and the distal half holotype is a essentially a pale, translucent fish, contains scattered melanophores and red and yel- with the anterior region of the dorsum suffused low chromatophores. The caudal fin contains a mix with light yellow, the exposed portions of the scales of yellow and red suffusions with scattered below the dorsal fins are thinly edged with red, melanophores, and the bases of the rays and the there are two yellow saddles a little less than pupil- regions adjacent to them are red, forming a thin diameter in width over the dorsal part of the red vertical line over the ends of the hypurals. The peduncle, and one (below the second dorsal saddle) basal region of the anal fin has a series of red over the ventral part of the peduncle. The neural inverted triangle separated by pale areas (these do canal and upper half of the swimbladder are heav- not conform to the bases of the fin elements), the ily invested with melanophores and clearly visible fin has a basal black stripe followed by a broad yel- in lateral view; the abdomen is pale pink anteriorly low stripe and culminating in a distal black stripe. grading to yellow posteriorly, the yellow intensify- The pectoral fin rays are red, intensified at their ing into a narrow, tapering wedge posteriorly above 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. 3 - 10 July 2011 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 Trimma haimassum n. sp. diffuse black margin and a horizontal, one-third (Figs 1 B & C, 2A, 6, 16-20) pupil-diameter dark purple stripe which touches Blood-spot pygmy goby the dorsal rim of the pupil ventrally. The second photographed specimen (17.3 mm SL, ROM Trimma RW sp. 63 – Dimara et al. 2010: 621 T7710, tissue voucher) is essentially similar, but (Raja Ampat) lacks the yellow wedge from the abdomen that Trimma sp. – Hayashi & Shiratori 2003: 44 (# passes posteriorly above the base of the anal fin. 070), 45 (# 078); Indonesia) Colour pattern in alcohol: off white, Trimma sp. 1 – Kimura & Matsuura 2003: 193 with darker pigmentation over the swimbladder (Bitung, Indonesia) visible through the body, as well as that covering Trimma sp. 1 (R. Winterbottom sp. 63) – Allen & the braincase around the posterior rim of the orbit. Erdmann 2009: 619 (Fakfak/Kaimana and Raja The dark pigmentation in the distal portions of the Ampat, Indonesia) dorsal fins, the base and distal regions of the anal Trimma sp. 10 – Kuiter & Tonozuka 2004: 706 fin, and a few dark chromatophores on the dorsal (Indonesia) surface of the snout are also visible. Comparisons: Trimma habrum belongs to the T. Material Examined: A total of 11 collections, 240 tevegae , which is defined by the (10.4-30.9 mm SL) type specimens, plus addi- apparently apomorphic conditions of the broad tional non-type specimens from Raja Ampat bony interorbital (75% or more of pupil diameter) (including a tissue voucher specimen), as well as and a Type A abdominal/caudal vertebral transi- numerous specimens from several other western tion region. Among the 10 nominal species cur- Pacific localities. The morphological description is rently assigned to this group, T. habrum differs based primarily on 10 males, 9 females (25.0-30.1 from T. caudomaculatum Yoshino & Araga, 1975, mm SL, –x = 27.6, SD = 1.52, ROM 85347), plus T. griffithsi Winterbottom, 1984, T. nasa Winter- the holotype, with some details (teeth, vertebral bottom, 2005 and T. tevegae Cohen & Davis, transition) from ROM 1836CS (5, 19.5-27.3); all 1969, as well as from T. xanthochrum n. sp. specimens from the same collection. described herein, in lacking any dark pigmentation Holotype: ROM 87484, 27.3 mm SL female, E on the posterior portion of the caudal peduncle. side of “Barracuda Rock”, 200 m N of Wayil Island All the remaining species except T. marinae Win- (02°11’43.4”S, 130°25’37.9”E), cave with lots of terbottom, 2005, have at least some branched rays sea fans & sea whips near entrance at top part, big in the middle of the pectoral fin. Trimma marinae variety of soft & hard corals just outside, 19.8-27.4 shares with T. habrum the thin vertical red bar over m, rotenone, 1300-1340, field # RW10-40, 1 Feb- the ends of the hypurals in freshly collected speci- ruary 2010, R. Winterbottom, L. Katz & CI team. mens, but has an open nasal capsule lacking ante- Paratypes: AMS I.45593-001 (formerly ROM rior and posterior nares (vs. a nasal sac with both 85390), 38, 14.6-30.9, SE islands off Misool, nares present), fewer scales in the predorsal midline south side of Balbulol Island (02°01’29.5”S, (6-8 vs. 9-10), and the bases of the elements of the 130°41’34.9”E), 45 m, rotenone, 0750-0810, field dorsal fin are not surrounded by a small, dark red # RW10-45, 2 February 2010, M. V. Erdmann. spot (vs. such a spot present). MZB 19779 (formerly ROM 85377), 59, 14.1- Distribution: Trimma habrum is currently only 27.7, SE islands off Misool, south side of Balbulol known from a single collection at 70 m on a verti- Island (02°01’29.5”S, 130°41’34.9”E), sea fans, cal wall at Keruo Island (off Penemu Island), one of sea whips, sponges, tunicates, hydroids, 13.7-18.3 the Fam Islands in Raja Ampat, Indonesia (Fig. 6). m, rotenone, 0750-0850, field # RW10-44, 2 Feb- Etymology: Derived from the Greek word ruary 2010, R. Winterbottom, L. Katz & CI team. ‘habros’, meaning delicate, dainty or graceful, in ROM 84877, 5, 21.0-27.5, Keruo Island, off Pen- allusion to the soft and delicate shades of colour of emu Island, Fam Islands (00°35’15.4”S, freshly collected specimens. Trimma habrum has 130°17’41.1”E), various hard and soft corals, tuni- been referred to informally (in litt.) as T. RW sp. cates and sponges, 18.3-27.4 m, clove oil, 0810- 95. 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 aqua vol. 17 no. 3 - 10 July 2011 146 Richard Winterbottom

(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, Kawe Island, SW bay (00°05’15.2”S, 130°07’25.3”E), 50 m, clove oil, field # RW10-10, 26 January 2010, M. V. Erdmann. ROM 85084, 1, 15.5, about 1.2 km SSE of Mutus Island, on west side of sand spit (00°21’01.0”S, 130°21’25.4:E), sponges, tunicates, some hard and soft coral, 50 m, clove oil, Fig. 16. Left lateral view of accessory scales above end of 0900-0920 field # RW10-16, 28 January 2010, M. pectoral fin of Trimma haimassum (27.3 female paratype, V. Erdmann. ROM 85156, 20, 14.3-25.8, Tanjung ROM 85347). Specimen stained with cyanine blue. Photo by R. Winterbottom. Manare, Waigeo Island, west side at about middle of 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. ROM 87415, 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 February 2010, M. V. Erdmann. ROM 1836CS, 5, 19.5- 27.3, collected with the holotype. USNM 399287 (formerly ROM 85210), 35, 10.4-24.9, Wofoh Island, west coast near south end (00°15’21.9”S, 130°17’32.0”E), many small corals, Tubastrea, sea whips, 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, ROM T07712 (24), collected with ROM 84884, for genetic 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); ROM 85106, 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); ROM 87452, 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, Fig. 17A-B. Trimma haimassum, 27.3 female paratype, WAM P.28161-011, 5 (12-24). Hermit Is., Jalun ROM 85347, Wayil Island, Raja Ampat. Left lateral (A) Island, USNM 264557, (22); Madehas Island, and dorsal (B) views of head to show papillae, specimen WAM P.28165-011, (18). Manus Island, ex-WAM stained with cyanine blue. Photo by R. Winterbottom. P.27826-086, (18). New Britain, Rabaul, ex-WAM

147 aqua vol. 17 no. 3 - 10 July 2011 Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature

P.28172-041, (25); WAM P.28174-025, 5 (17-29). about 55% the width of the pupil and with a fleshy Philippines: Cebu, ROM 1137CS, 3 (19-30); median ridge between the eyes, and a dermal ridge ROM 54753, 8 (19-30). Negros Oriental, ROM anterior to the first dorsal spine. The red spot 54742, 6 (22-26). Palawan, El Nido, BLIH above the opercle, which continues a thin red line 1983354, 3 (25-28). Siquijor Island, ROM 54741, below the orbit where it is underlain by a blue (23); ROM 69534, (24); ROM 69583, (17). stripe, is distinctive in live individuals. In fresh and Solomon Islands: Guadalcanal, ROM 54754, preserved material, the red spot and line are dark, (23); ROM 54755, 2 (15-22). there are two dark stripes between the anteroven- Diagnosis: Trimma haimassum is characterized by tral margin of the eye and the edge of the upper lip, a relatively deep body, numerous irregular scales and the tip of the snout has a dark reticulated pat- with about 30 lateral rows and over 15 scales in the tern, with dark transverse stripes over the dorsal anterior transverse series, a fifth pelvic fin ray that margin of the orbit. branches twice dichotomously, usually a somewhat Description: The number of specimens (n) is elongate second dorsal spine reaching to a little given only where they are less than 20. The body beyond the spine of the second dorsal fin, no scales scales in the species are in numerous, irregular rows on the midline of the predorsal region, cheek or that may be partially incomplete, and accessory opercle, a moderately wide bony interorbital that is scales are usually present (Fig. 16), making scale

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. aqua vol. 17 no. 3 - 10 July 2011 148 Richard Winterbottom counts exceedingly difficult and potentially unreli- 2 unbranched (n = 7), once 7/2. Lateral scales able. Dorsal fins VI + I 10, second spine longest, about 29-31 (x– = 30.3, SD = 0.63), anterior trans- usually reaching to between bases of first to fourth verse scales about 15-16-17 (x– = 16.7, SD = 0.83), rays of second dorsal fin when adpressed (not elon- posterior transverse scales about 10-12-13(x– = gate in two males, 29.6-29.9 mm SL, reaching 11.6, SD = 0.79), no scales in midline of predorsal, either to base of spine of second dorsal fin or inter- cheek or on opercle; pectoral base with 4-5 vertical space between fins), all rays branched except, usu- rows of cycloid scales, outer row of 7-8-10 (n = 16) ally, posterior element of last ray; anal fin I 8-9-10 scales, anteriormost row largest, posteriormost (x– = 9.0, SD = 0.37), all but first and posterior ele- smallest; 7-9 prepelvic cycloid scales in midline ment of posteriormost ray usually branched (first anterior to pelvic-fin base (2 specimens with 11, element branched in four specimens); posterior- where scales extended anterior to ventral tips of most ray of both fins reaching between half and cleithra on isthmus); 16-18 circumpeduncular three-quarters of distance between ray base and scales (x– = 16.9, SD = 0.70, n = 18); body scales first dorsal or ventral procurrent ray; pectoral fin ctenoid, extending from above posterior third of 18-19 (x– = 18.5, SD = 0.50), 3-4 (x– = 3.2, SD = opercle, passing slightly anteriorly and then up and 0.39) and 1-3-4 (x– = 2.5, SD = 0.66) unbranched back in an arc, with a narrow scaleless border along dorsal and ventral rays respectively, middle rays base of first dorsal fin up to about base of 5th ray; branched, fin reaching posteriorly to a vertical line scales cycloid anterior to a line about 3 scales pos- between middle of urogenital papilla and base of terior to axil of middle of pectoral base to about anal spine; pelvic fin I 5, no frenum, basal mem- mid region of belly. Gill opening extending brane 25-55% length of fourth ray (n = 15, high anteroventrally to a vertical below anterior third to variance probably due to delicate nature of mem- middle of pupil. Upper jaw with outer row of brane, which is easily torn), first four rays with 2-4 spaced, curved, enlarged canines, decreasing some- sequential branch points, fifth ray branched what in size posteriorly and extending to end of branched twice dichotomously and 59-68-76% premaxilla, 3-4 irregular rows of small conical teeth length of fourth (x– = 67.6, SD = 4.51), fourth ray behind the outer row at symphysis, those near sym- reaching posteriorly to between urogenital papilla physis in innermost row directed posteriorly, and base of second anal fin ray. Although seg- enlarged, but decreasing to size of other inner row mented caudal fin rays nearly always total 17 (1 of teeth posterolaterally, which grade to single row 18 specimens with 16), variation in number of posteriorly at end of premaxilla. Lower jaw with branched vs. unbranched rays was found; dorsal outer row of spaced, curved, enlarged canines, end- half usually with 3 unbranched and 6 branched ing at bend of dentary where these teeth are largest, rays (once 2/6, once 1/7); ventral half usually with several irregular rows of small conical teeth at sym- 5 branched, 3 unbranched (n = 10) or 6 branched, physis grading to single outer row ending at

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

149 aqua vol. 17 no. 3 - 10 July 2011 Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature anterodorsal tip of coronoid process of dentary, distal to the base, and the fin membranes are heav- innermost row somewhat enlarged (half the size of ily sprinkled with melanophores. The caudal fin outer teeth), spaced, and continuing posteriorly to has scattered, half-pupil diameter sized yellow to beginning of coronoid process. Cephalic sensory orange spots and numerous melanophores in the papillae as in Fig. 17 and Table I (nomenclature of fin membranes. The anal fin also has numerous papillae given in Fig. 2A). Tongue truncate with melanophores in the membranes, and is suffused rounded edges. Gill rakers on first arch 4-5 (x– = with reddish-pink. The pelvic fin is pinkish, the 4.4, SD = 0.49) + 16-17-18 (x– = 16.9, SD = 0.71) pectoral fin has reddish fin rays and hyaline mem- = 20-21-22 (x– = 21.3, SD = 0.71), larger rakers on branes. A 27.5 mm SL male (Fig. 19) has a more both epibranchial and ceratobranchial possess a orange head and body, the blue lines under and in row of small, bluntly pointed crenulations along front of the eye are intermittent or absent as is the their medial surfaces (Fig. 1B). Anterior nasal red stripe between the ovoid spot above the end of opening a short tube extending out over upper lip, the opercle and the opercle, the basal stripe in the posterior nasal opening a pore with a raised rim, dorsal fins is vivid orange, there is a line of orange both protruding from slightly raised oval sac con- spots in the second dorsal at about the middle of fined to anterior half of snout, posterior opening their length, the caudal spots are orange, as is the about 2-3 X its diameter from closest border of eye. basal half of the anal fin, and the scale pockets on Bony interorbital 47-64% pupil width (x– = 55.5, the dorsal half of the body are more heavily out- SD = 5.07), interorbital profile broadly U-shaped lined with dark chromatophores. A specimen pho- with a central ridge of soft tissue separating the left tographed underwater (Fig. 20) has a much yel- and right sensory papillae row p (Fig. 17 B). Ridge lower head, the red stripe from the ovoid opercular of tissue (dermal crest) extending from proximal spot is in two parts, and the bars across the orbit one-tenth of first dorsal spine anteriorly, becoming and lines on the snout are blue. An anaethetised reduced in height anteriorly and fading out above photographed underwater has the red opercular anterior third opercle. Epaxialis musculature blotch continuing anteriorly across the ventral extending anteriorly to above a vertical with poste- margin of the pupil almost to the maxilla, and the rior third of pupil. Abdominal/caudal vertebrae blue line beneath it extends posteriorly to the mid- transition Type B. region of the opercle. The area immediately above Colour pattern (based on images of an this red stripe on the snout is bright yellow, and the anaesthetised specimen and nine freshly collected thin blue stripe above it is margined with red. The specimens). A 25.9 mm SL gravid female (Fig. 18) diagonal stripe across the iris is mauve, as are the has a reddish head, orange-red dorsum grading to short stripes to the midline on the fleshy part of the yellowish pink posteriorly, slightly lighter over the anterodorsal orbit. The median fins are more abdomen, with some of the more dorsal scale mar- greenish-yellow, but this may be due to the dark gins slightly darker. An ovoid red spot, about blue background, and the scale pockets on the dor- pupil-diameter in width lies on and above the pos- sum are relatively strongly outlined. The remaining terodorsal margin of the opercle and is connected photographed specimens are similar overall, to the posteroventral margin of the orbit by an although some have many more dark pigment cells intermittent red stripe. A thin blue line underlies in the fin membranes (including the pelvic fin), the anterior portion of this stripe and continues may have a more or less distinct dark basal band in anteroventrally, skirting the ventral margin of the the dorsal fins, and the overall body colour may be orbit and ends just above the upper jaw. A short brown rather than red, pink or yellow. blue line from the orbit to just above the middle Colour pattern in alcohol: head and length of the maxilla is present just dorsal to this body pale straw-yellow; the nape, upper half of the line, and is similarly oriented. Four short blue lines opercle and the body, except for the anteroventral between the anterior and dorsal margins of the region of the abdomen, with a dense scattering of fleshy orbit pass medially towards the midline. The melanophores and subdermal brown chro- iris is red and speckled with darker pigment, and matophores, which tend to be concentrated around there is a thin blue line passing across it from the scale pockets anterodorsally in specimens in anteroventral to posterodorsal oriented at about which the scales have been abraded off. An ill- 45°. The dorsal fins have an orange stripe, about defined, irregular line primarily of mela no phores one-third pupil diameter in width, situated just runs anteroventrally from the anterodorsal region of aqua vol. 17 no. 3 - 10 July 2011 150 Richard Winterbottom the opercle across the ventral margin of the orbit and opercular margin anteriorly to the posterior margin ends above the maxilla; a similar short stripe lies of the eye (vs. absent). above this and passes parallel to it from the Distribution: Trimma haimassum has been col- anteroventral margin of the orbit to above the mid- lected from south-western Sulawesi north to dle of the maxilla. The snout and both lips have an Palawan in the Philippines and eastwards to the irregular reticulated mottling of melano phores. The Solomon Islands (Fig. 6) in depths of 7-70 m. It dorsal margin of the orbit has four to five transverse may also occur in Palau. This was the most abun- stripes of melanophores. The cheek and pectoral fin dant species of the genus at Raja Ampat. base have a few scattered melanophores, and the Etymology: Derived from the Greek word dorsal margin of the pectoral fin base occasionally ‘haimasso, stained with blood, in allusion to the has a thin dark stripe; the membranes of the blood-red spot above the posterior end of the oper- unpaired fins are heavily invested with melano - cle in live and fresh specimens. This species has phores, while those of the paired fins have only a few been referred to informally (in litt.) as Trimma RW widely scattered melano phores. There is little evi- sp. 63. dence of the sub-basal stripe in the dorsal fins pre- sent in freshly collected specimens, although some Trimma papayum n. sp. specimens do seem to have fewer melano phores in (Figs. 6, 21-24) the region where the stripe occurs. Pawpaw pygmy goby Comparisons: The new species belongs to the T. sheppardi species complex, a group which lacks Trimma RW sp. 93 – Dimara et al. 2010: 621 median predorsal, cheek and opercular scales, pos- (Raja Ampat) sesses a deep head and body (body depth at base of Trimma sp 6 – Kuiter & Tonozuki 2004: 705 (two pelvic spine about 30% SL), has accessory scales underwater photographs, Maumere, Flores) and disjunct scale rows, has a well developed nasal apparatus occupying the anterior half of the snout Material Examined: A total of 2 lots, 12 type spec- with the posterior opening in a short tube, has imens, plus one additional non-type specimen (tis- median fleshy ridges in the interorbital region and sue voucher). The description is based on the holo- possesses a dermal crest on the nape just anterior to type and up to 11 paratypes (11.5-22.2 mm SL). the origin of the first dorsal fin. Trimma haimassum Holotype: ROM 87485, 21.6 mm SL female, differs from all the other species in the group by Indonesia, Raja Ampat, SW bay of Kawe Island, nearly always having an elongate second dorsal fin 00°05’15.2”S, 130°07’25.3”E, 50 m, clove oil, 26 spine (vs. rarely elongate), having two (vs. one) January 2010, M. V. Erdmann. dichotomous branch points in the fifth pelvic fin Paratypes: AMS I.45594-001, (20.5), collected ray, in apparently in having more lateral scale rows with the holotype. MZB 19780, 4 (16.8-20.5), (29-31 vs. 24-26 – uncertainty due to the difficulty collected with ROM 87564. ROM 85034, 2, in accurately recording this count), in the dark 12.2-21.1 mm SL, collected with the holotype. reticulations on the dorsal surfaces of the snout and ROM 1833CS, 19.7 mm SL, collected with the lips, in having two dark stripes between the eye and holotype. ROM 87564, 3 (11.5-22.2), Indonesia, the maxilla, and in the colouration of the dark spot Raja Ampat, SW Kawe Island, 00°05’46.6”S, 130° above the opercle in life (red vs. black). It differs 07’07.2”E, 50 m, clove oil. M. V. Erdmann, 28 further from T. sheppardi Winterbottom, 1984, in August 2010. having 10 (vs. 8-9) dorsal-fin rays; more outer gill Additional (Non-type) Material. One specimen rakers on the first gill arch (usually 4-5 + 15-18 vs. for genetic analysis, ROM T07725, (20.5), col- 2-3 + 13-15); in lacking the two yellow bars on the lected with the holotype. cheek beneath the eye; and in having the anterior Diagnosis: Trimma papayum is unique among the portion of the head orange-red rather than pink. It described species of the genus in having a one-third differs from T. yanoi Suzuki and Senou, 2008, in pupil diameter black spot on and just behind the lacking the obvious internal dark blotches along fourth dorsal fin spine. It has 9 dorsal and 8 anal the vertebral column that are especially prominent fin rays, 18-19 pectoral rays with the middle ones in live and freshly collected material. It differs fur- branched, a single branch point in the fifth pelvic ther from an undescribed species (T. RW sp. 24) in fin ray, which is about 60-70% the length of the having the extension of the dark areas above the fourth, and a basal membrane about 10-20% the

151 aqua vol. 17 no. 3 - 10 July 2011 Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature length of the fourth ray, 10-11 anterior and 8-9 of 3 scales on the upper margin of the opercle, and posterior transverse scale rows, 5-10 scales in the a shallow interorbital trench with no trench pos- predorsal midline, no cheek scales and a single row terodorsal to the orbit. Freshly collected specimens are orange red in overall colouration, with scattered diffuse yellow spots. Description: Dorsal fins VI + I 9, second and third spines longest, second reaching to between base of fifth ray of second dorsal fin and two scales beyond end of that fin when adpressed, third reaching to bases of first to fourth ray of second dorsal 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 dorsal procurrent fin ray (x– = 67.1, SD = 10.10, n = 12) ; anal fin I 8, all but first and posterior element of posteriormost ray branched; posteriormost ray reaching between 50-75% (x– = 58.7, SD = 7.59, n = 12) of distance from its base to first ventral procurrent fin ray; pectoral fin 18-19 (x– = 18.4, SD = 0.49, n = 12), 3-4 dorsal and 3-6 ventral unbranched rays (but 6 and 9 respectively in the 11.5 mm SL juvenile), middle rays branched, fin reaching posteriorly to between a vertical line above urogenital papilla and base of first ray of anal fin; pelvic fin I 5, no frenum, basal membrane 9- 18% length of fifth ray, first four rays with one sequential branch (once with two branch points in fourth ray), fifth ray branched once dichotomously and 56-65-74% (x– = 64.8, SD = 5.15, n = 10) length of fourth, fourth ray reaching posteriorly to between bases of first to fifth anal-fin rays. Lateral scales 23 (n = 11); anterior transverse scales 9-11 (x– = 10.1, SD = 0.69, n = 11); posterior transverse scales 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 11.5 mm SL specimen); no scales on cheek; a single row of 3 cycloid or weakly ctenoid scales on opercle; 3-4 vertical rows of scales on pectoral fin base with 4-5 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 midline between base of last anal ray and first procurrent caudal ray; body scales ctenoid except for cycloid scales on anterior belly midline, breast, beneath and just posterior to pectoral-fin; body scales extend anteriorly to vertical in line with posterior margin of orbit. Gill opening extending anteroven- Fig. 21A-C. Left lateral (A), dorsal (B), and ventral (C) trally to a vertical below middle to anterior one- views of head to show head papillae of Trimma papayum third of pupil. Upper jaw with outer row of slightly (19.6 mm SL male paratype, ROM 87564). Specimen curved, enlarged canines decreasing in size posteri- stained with cyanine blue. Photo by R. Winterbottom. orly to end of premaxilla, 2-3 irregular inner rows aqua vol. 17 no. 3 - 10 July 2011 152 Richard Winterbottom

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

almost to a vertical above posterior margin of orbit. Abdominal/caudal vertebral transition Type B. Colour pattern (from a freshly collected specimen, 21.6mm SL female holotype, Fig. 22). Overall orange-red (red especially evident along the middle of the trunk, above the anal fin and on the peduncle), with numerous diffuse pupil-diam- eter yellow spots on the body at the centres of the scales, decreasing somewhat in size both anteriorly on the nape and posteriorly on the peduncle, those Fig. 23. Left lateral view of the first dorsal fin to show the on the nape yellow-orange rimmed with red, the size and position of the dark ocellated spot of Trimma rest of the nape and the area immediately below the papayum (21.1 mm SL female paratype - preserved, ROM dorsal fins grey with the exposed margins of the 85034). Note damaged second dorsal spine. Photo by R. scales diffusely rimmed with dark chromatophores; Winterbottom. the belly is pale. The snout is darkened with a mix- of small conical teeth near symphysis decreasing to ture of chromatophores and melanophores; the a single row to end of premaxilla. Lower jaw as for cheek is red with a slight yellow suffusion anteri- upper jaw, but outer row ends at bend of dentary, orly, two faint yellow spots below the anteroventral inner row enlarged at symphysis and decreases in margin of the eye; the ventral region of the cheek size posteriorly to end at coronoid process. and the branchiostegal rays are a darker red. The Cephalic sensory papillae as in Fig. 21 and Table I. pectoral fin base has two reddish-orange spots, the Tongue truncate with rounded margins. Gill rakers dorsal of which lies a little anterior to the ventral, on first arch 4-5 (x– = 4.4, SD = 0.48) + 14-16 (x– = which latter is the darker of the two. The first dor- 14.6, SD = 0.77) = 18-20 (x– = 19.0, SD = 0.95, n sal fin has a thin yellow-orange stripe just distal to = 11). Anterior nasal opening a short broad tube the base between the first three spines. The anterior extending out over upper lip, posterior nasal open- margin of a small (one-third pupil-diameter) ovoid ing a pore with a raised rim, both protruding from black spot overlies, or may begin just posterior to, slightly raised oval sac, posterior pore about 1 pore- the fourth dorsal spine and is confined to the diameter from nearest point of bony orbital rim. region between the fourth and fifth spines. It lies Bony interorbital 22-40-47 (x– = 33.7, SD = 6.91) on the same plane as the stripe (Fig. 23), and has a % pupil width, with broad shallow U-shaped wide (about equal in width to the black spot) red interorbital trough and no postorbital trough or halo except dorsally over the spine, where the halo trench. No or very slight dermal crest anterior to grades to yellow. A live specimen photographed at first dorsal fin, epaxialis musculature reaching 50 m is generally similar, but has a darker body, a

153 aqua vol. 17 no. 3 - 10 July 2011 Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature faint thin light stripe beneath the eye, light vermic- tral, and two spokes of a similar size and colour pass ulations over the snout and interorbital region, two dorsally towards the midline from the upper surface pupil diameter orange spots on the opercle, and the of the pupil. red halo around the dark spot in the dorsal fin lacks Colour pattern in alcohol: As above, the yellow dorsal margin. A tiny black spot, situated but pale straw coloured (all red, orange and yellow on the first dorsal spine just above its base, is present areas pale), dark pigmentation outlining of scale in two of the 12 specimens, a 12.1 mm SL juvenile pockets and in fin membranes remaining. and an 18.9 mm SL female. A similar tiny black spot Comparisons: Trimma papayum belongs to a is present on and behind the second spine in a pho- grade of 24 valid described species of Trimma, all tographed specimen from Maumere, Flores. The of which have a Type B vertebral transition pattern second to sixth spines have elongate yellow spots a and scales present in the predorsal midline. In 11 little distal to the stripe and spot, and the fin mem- of these species, the second dorsal spine reaches no branes are invested with dark chromatophores and further posteriorly than the spine of the second melanophores, especially proximally and distally. dorsal fin when the fin is adpressed (reaches poste- The second dorsal fin has numerous yellow spots, riorly to between the base of the fifth dorsal ray to the proximal (just above the base of the fin) series beyond the last ray in T. papayum). Four of the forming an irregular, interrupted stripe, the others remaining species, T. lantana Winterbottom & less obviously forming stripes, and the fin mem- Villa, 2003, T. mendelssohni (Goren, 1978), branes contain numerous dark chromatophores and T. naudei Smith, 1957 and T. okinawae (Aoyagi, melanophores with a reddish suffusion at the level of 1949) have a deep postorbital trench (vs. a trench the proximal row of yellow spots. The anal and cau- absent in T. papayum). Trimma randalli Winter- dal fins are similar, with scattered yellow spots, a red- bottom & Zur, 2007, has an unbranched pectoral dish suffusion proximally, and numerous dark chro- fin ray (vs. several middle rays branched) and matophores and melanophores (especially distally). T. flavicaudatum (Goren, 1982) and T. yanagitai The membranes of the pectoral and pelvic fins are Suzuki & Senou, 2007, have 8 or fewer dorsal fin hyaline, with the fin rays of the former reddish and rays (vs. 9). Of the six remaining species, T. barralli those of the pelvic fin yellowish. The dorsal three- Winterbottom, 1995, has more lateral scales (25- quarters of the iris is dark red, with the ventral area 26 vs. 23) and is confined to the Red Sea, T. imaii being red and yellow. A thin charcoal to dark purple Suzuki & Senou, 2009, has 16 pectoral fin rays (vs. line bisects the iris from anterodorsal to posteroven- 18-19), T. nomurai Suzuki & Senou, 2007, has a

Fig. 24. Trimma papayum (live), Kawe Island, Raja Ampat. Photo by M. V. Erdmann. aqua vol. 17 no. 3 - 10 July 2011 154 Richard Winterbottom pupil-diameter dark round spot on the body soft coral, 50 m, clove oil, 0900-0920, field # between the pectoral and first dorsal fin (vs. such a RW10-16, 28 January 2010, M. V. Erdmann. spot absent), T. preclarum lacks scales on the oper- Paratypes: AMS I.45595-001 (formerly ROM cle (vs. present) and has fewer anterior and poste- 85362), 4, 20.7-23.4, E. side of “Barracuda Rock”, rior transverse scale rows (7-8 and 6-7 vs. 9-11 and 200 m N of Wayil Island (02°11’43.4”S, 8-9 respectively), and T. squamicana Winterbot- 130°25’37.9”E), 60 m, clove oil, 1300-1320, field tom, 2004, has an unbranched fifth pelvic fin ray # RW10-41, 1 February 2010, M. V. Erdmann. (vs. branched once) and lacks scales on the opercle MZB 19781 (was ROM 85145), 25, 16.5-21.3, (vs. present). Trimma halonevum Winterbottom, Jef Tsiep Island, west side, channel between it and 2000, is perhaps the most similar species to T. small island to the west (00°23’05.7”S, papayum in meristic values and in general shape, 130°16’37.1”E), 42 m, clove oil, 1600-1620, field but adults have 2-3 cycloid scales on the cheek. It # RW10-21, 20100128, M. V. Erdmann. ROM has a very similar small ocellated dark spot in the 84885, 4, 7.4-25.1, Keruo Island off Penemu first dorsal fin, but this is confined to the mem- Island, Fam Islands (00°35’15.4”S, 130°17’ brane between the fifth and sixth spines (vs. on and 41.1”E), 70 m, clove oil, 0815-0850, field # just posterior to the fourth spine), and has numer- RW10-04, 25 January 2010, M. V. Erdmann. ous similar dark haloed spots scattered over the ROM 84891, 2, 19.0-19.3, Keruo Island off Pen- body (vs. such body spots absent). emu Island, Fam Islands (00°35’15.6”S, 130°17’ Distribution: Trimma papayum (Fig. 6) is cur- 41.1”E), 56 m, clove oil, 1200-1300, field # rently known only from Maumere, Flores (photo RW10-05, 25 January 2010, M. V. Erdmann. only) and Kawe Island in the Raja Ampat region of ROM 85082, 8, 17.5-21.6, collected with the Indonesia, where it was collected at a depth of 50 m. holotype. ROM 85132, 1, 16.6, Jef Tsiep Island, 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 black spot or spots in the first dorsal fin characteristic of the new species. Trimma papayum has been referred to informally (in litt.) as Trimma RW sp. 93.

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

Trimma RW sp. 94 – Dimara et al. 2010: 621 (Raja Ampat) Trimma tevegae (Cohen & Davis, 1969) – Allen & Erdmann 2009: 619 (in part, Bird’s Head region, Indonesia)

Material Examined: A total of 13 collections, 123 type specimens (7.4-25.1 mm SL), plus 19 non- types (including 7 tissue voucher specimens). The description is based on up to 23 specimens from ROM 85082, 85155, 85208, 85225, 85319, 87483 and 1834CS (17.7-24.1, 14 males, 7 females, 2 unsexed). Holotype: ROM 87483, 22.3 mm SL male, Fig. 25A-B. Head in A. left lateral (reversed) and B. dorsal Indonesia, Raja Ampat, about 1.2 km SSE of Mutus view to show head papillae in Trimma xanthochrum (22.3 Island, on west side of sand spit (00°21’01.0”S, mm SL male holotype, ROM 87483). Specimen stained 130°21’25.4”E), sponges, tunicates, some hard and with cyanine blue. Photo by R. Winterbottom.

155 aqua vol. 17 no. 3 - 10 July 2011 Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature west side, channel between it and small island to 0750-0810, field # RW10-45, 2 February 2010, M. the west (00°23’05.7”S, 130°16’37.1”E), variety of V. Erdmann. ROM 1834CS, 5, 9.9-20.9, collected hard & soft corals, sponges, 19.8-22.9 m, clove oil, with the holotype. USNM 399288 (formerly ROM 1600-1630, field # RW10-20, 28 January 2010, R. 85190), 3, 18.8-20.4, Tanjung Manare, Waigeo Winterbottom, L. Katz & P. Johannes. ROM Island, west side at about middle of width off small 85155, 13, 9.5-21.2, Tanjung Manare, Waigeo cape (00°16’26.1”S, 130°19’01.5”E), sponges, hard Island, west side at about middle of width off a corals, 52 m, rotenone, 0750-0810, field # RW10- small cape (00°16’26.1”S, 130°19’01.5”E), hard & 24, 29 January 2010, M. V. Erdmann. soft corals, sponges, 19.8-22.9 m, rotenone, 0750- Additional (Non-type) Material. Seven specimens 0820, field # RW10-23, 29 January 2010, R. Win- for genetic analysis: one collected with ROM 84885 terbottom, L. Katz, P. Johannes, W. Kaka & W. (ROM T07708, 25.2); two with ROM 85082 Awom. ROM 85208, 2, 18.5-20.2, Wofoh Island, (ROM T07735, 20.7, T07736, 21.8), two with west coast near south end (00°15’21.9”S, ROM 85155 (ROM T07748, T07749, 21,8), one 130°17’32.0”E), many small corals, Tubastrea, sea with ROM 85208 (ROM T07752, 23.2) and one whips, some black coral, sea fans, tunicates, with ROM 85319 (ROM T07761, 24.8). ROM sponges, some hydroids, 18.3-22.9 m, clove oil, 84934, 12 (9.9-12.5), collected with the holotype). 1500-1540, field # RW10-25, 29 January 2010, R. Diagnosis: Trimma xanthochrum is characterized Winterbottom & W. Kaka. ROM 85225, 21, by a wide interorbital region (80-100% pupil diam- 15.2-22.0, Wofoh Island, west coast near south eter), a second dorsal spine reaching posteriorly to end (00°15’21.9”S, 130°17’32.0”E), many small between the bases of the spine of the second dorsal corals, Tubastrea, sea whips, some black coral, sea fin and the fifth rays, 15-16 pectoral rays of which fans, tunicates, sponges, some hydroids, 12-16 m, the middle 4-9 (usually 7-8) are branched, vertical clove oil, 1610-1700, field # RW10-27, 19 January rows of 2-3 sensory papillae below eye in rows 1-4 2010, L. Katz. ROM 85319, 3, 22.3-24.1, Kepot- and 4-5 papillae in row 5, a caudal spot which has a sol Island, north side at about middle of length, SE upper half about two-thirds the width of the lower of Misool Island (02°09’32.1”S, 130°17’34.0”E), half, and usually an overall yellowish body with yel- huge variety of soft & hard corals, some sponges, low at least proximally in the caudal fin. tunicates, & hydroids, 18.3-21.3 m, clove oil, Description: Dorsal fins VI + I 7-8 (once 7, n = 0830-0900, field # RW10-37, 21 January 2010, R. 23), second and third spines longest, second reach- Winterbottom, L. Katz & CI team. ROM 85333, ing to between base of spine to fifth ray of second 17, 15.0-24.5, Kepotsol Island, east side dorsal fin (usually to between bases of second and (02°09’32.1S, 130°17’34.0”E), 66 m, clove oil, third rays), third reaching to between just anterior 0830-0850, field # RW10-38, 1 February 2010, M. to dorsal spine to base of first ray of second dorsal V. Erdmann. ROM 85389, 14, 16.7-25.0, SE fin, rays all branched except posterior element of islands off Misool, south side of Balbulol Island last ray, last ray reaching to 35-45-50% distance (02°01’29.5”S, 130°41’34.9”E), 45 m, rotenone, between its base and first dorsal procurrent caudal

Fig. 26. Left lateral view of Trimma xanthochrum, 21.4 mm SL male paratype, ROM 85082, Mutus Island, Raja Ampat. Photo by R. Winterbottom. aqua vol. 17 no. 3 - 10 July 2011 156 Richard Winterbottom ray (x– = 42.0%, SD = 4.19, n = 20); anal fin I 8, SD = 4.07, n = 20); pectoral fin 15-16 (x– = 15.8, first ray and posterior element of last ray in 17 (of SD = 0.38, n = 23), 3-4-9 dorsal and 3-4-6 ventral 23) unbranched; posteriormost ray reaching unbranched rays respectively (more unbranched between 28-35-43% distance between its base and rays in smaller specimens), middle rays branched, first ventral procurrent caudal ray (x– = 35.4%, fin reaching posteriorly to a vertical line between posterior 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 of fourth (x– = 51.1%, SD = 3.78. n = 18), fourth ray reaching posteriorly to between bases of anal spine to fourth anal-fin ray. Lateral scales 23-24 (x– = 23.2, SD = 0.36, n = 20), when 24, first scale in pectoral fin axil is half the size of next scale and cycloid; anterior transverse scales 9; posterior trans- verse scales 8; 12-14 irregular scale rows in midline of predorsal (x– = 12.8, SD = 0.54, n = 20); three rows of cycloid scales on cheek, dorsalmost row of 3-4, 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 on opercle, with 1-3-4, 3-4, 2-4 and 1-2 cycloid, mid- dle scales of second row may be ctenoid, once a fifth row of a single cycloid scale; 3 vertical rows of cycloid scales on pectoral base with 4 scales (once 3) 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; 9-10 scales in midline between base of last anal ray and first ventral procurrent ray (x– = 9.9, SD = 0.31, n Fig. 27A-B. Left lateral view of the caudal peduncle to = 19); body scales ctenoid except for cycloid scales show differences in caudal spot of A. Trimma xanthochrum on anterior belly midline, beneath and just poste- (20.9 female, ROM 85155) and B. T. tevegae (21.0 female, rior to pectoral-fin base, and one or two scales adja- ROM 85320). Photo by R. Winterbottom. cent to posterodorsal margin of orbit. Gill opening

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

157 aqua vol. 17 no. 3 - 10 July 2011 Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature extending anteroventrally to a vertical below poste- anteroventral and posteroventral margins of eye. rior one-third of pupil. Upper jaw teeth of a row of Papillae immediately below eye on cheek in five enlarged spaced, curved caniform teeth that transverse rows, usually with 2, 2, 3, 3 and 5 papil- decrease about 50% in size from symphysis to dis- lae in vertical rows 1 through 5 (Fig. 25A – see also tal tip of premaxilla, two to three inner rows of Fig. 2B). Tongue truncate with rounded edges and smaller conical teeth at symphysis, innermost row a shallow V-shaped tip. Gill rakers on first arch slightly larger and curved, grading to a single row 3-4 + 14-17 = 18-21 (x– = 3.9 + 15.5 = 19.4, SD = posteriorly to end at distal tip of premaxilla. Lower 0.28, 0.72, 0.87 respectively; n = 23). Anterior jaw teeth with an outer row of 6-8 enlarged curved, nasal opening a broad tube extending almost to spaced canines ending at bend of dentary, about above anterior margin of upper lip, posterior nasal four irregular rows of small conical teeth at symph- opening a pore with a raised rim, both protruding ysis, innermost row increasing in size posteriorly from slightly raised oval sac, nasal apparatus con- until almost as big as outer symphysial row at mid- fined to anterior one-third of snout. Bony interor- dentary, then gradually decreasing in size and end- bital 80-89-100% pupil width (mean = 91.4%, SD ing at dorsal tip of coronoid process. Sensory papil- = 6.67, n = 19), interorbital shallowly concave with lae very delicate and subject to abrasion, especially a broad, rounded median fleshy ridge and no pos- those on cheek, opercle and immediately above torbital trough or trench (Fig. 25B). No dermal opercle and posterior to eye, several specimens with crest anterior to first dorsal fin. Epaxialis muscula- supernumary papillae (not included in counts in ture extending anteriorly to above posterior margin Table I) which were most frequently found around of pupil in dorsal midline. Vertebral transition a

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

Fig. 30. Left lateral view of Trimma xanthochrum, 23.2 mm SL male paratype, ROM 85319, Kepotsil Island, Raja Ampat. Photo by R. Winterbottom. aqua vol. 17 no. 3 - 10 July 2011 158 Richard Winterbottom modified Type A, with the tenth abdominal and where the entire scale pockets are heavily outlined first caudal vertebrae (only) with a small basal with dark pigment. A diffuse dark blotch covers haemal canal, the main haemal arch of the first the hypural region and bases of the caudal rays and caudal vertebra expanded, that of the second about a thin, diffuse vertical yellow bar over the ends of half the size of the first, remaining caudal haemal the hypurals separates the blotch into anterior and arches normal; swimbladder terminates at the posterior halves, the posterior half being more haemal canal of second caudal vertebra. strongly pigmented than the anterior half. The Colour pattern (based on freshly collected dorsal half of the blotch is about two-thirds the material, a total of seven images, 4 males, 3 width of the ventral half (Fig. 27A), and does not females, and one live specimen). A 21.4 mm SL extend anteriorly beyond the base of the anterior- male (Fig. 26) has yellowish body (especially along most procurrent fin ray. The first dorsal fin has a the ventral region) with a diffuse rosy stripe above one-third pupil width dark basal stripe, followed the lateral septum to within about a scale width of by a pupil width yellow stripe, with the rest of the the dorsum. The lining of the swim bladder is fin heavily invested with melanophores; the spines darkly pigmented, which shows through the body are reddish. The second dorsal fin has a similar wall and is augmented by brown subdermal chro- dark basal stripe, above which a narrow yellow matophores over the upper half of the abdominal stripe expands posteriorly until it occupies the cavity, together forming a wedge-shaped darker entire height of the fin from the sixth fin ray on; stripe which fade out just posterior to the anal fin. the distal anterior portion of the fin has numerous The upper part of the head and upper jaw is heav- melano phores. The anal fin is essentially hyaline ily and darkly pigmented, with a thin diffuse red with a slight reddish suffusion on the posterior- stripe just below the eye from the vertical limb of most rays. The pectoral fin is yellow to orange with the preopercle to middle of the maxilla. The area a few scattered brown chromatophores except on below this stripe is light red with yellow suffusions, the dorsal rim, which is rimmed with melano- becoming pale ventrally. The body scales are thinly phores to form a diffuse, elongate spot; the fin is edged with melanophores along their exposed mar- hyaline with red fin rays. The pelvic fin is dusky. gins, except ventrally on the abdomen and trunk, The dorsal and ventral margins of the caudal fin where there is no such pigmentation, and dorsally, are hyaline with reddish unbranched caudal rays,

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.

159 aqua vol. 17 no. 3 - 10 July 2011 Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature the rest of the fin is yellow with a pale central prox- from all but T. caudomaculatum and T. tevegae in imal stripe and the membrane between the middle having a well developed large dark spot at the end of two rays has a red suffusion. The iris is densely mot- the caudal peduncle that extends the full width of tled with dark pigment, with a golden suffusion pos- the peduncle. In both those species, the caudal spot teroventral to the pupil; a dark stripe passes over the is rounded with both dorsal and ventral halves top of the pupil, above which is a diffuse reddish- approximately equal in diameter (Fig. 27B – vs. dor- yellow stripe. A 20.8 mm SL male (Fig. 28) differs sal half about two-thirds the width of the ventral in having the dorsal half of body pale, the thin red half – Fig. 27A), there are 12-14 unbranched pec- stripe below the eye is followed by a diffuse pale toral fin rays (vs. 15-16 rays with at least 4, and usu- stripe just below it, and the lower jaw, except the tip, ally 6-8, of the middle rays branched), the fifth is pale. The dorsal half of the caudal blotch is better pelvic ray is nearly always unbranched (vs. branched developed and about four-fifths the width of the once in 87% of specimens), and the vertical papillae ventral half; the proximal half of the caudal fin is in rows 2-4 just below the eye consist of a single much more intensely pigmented than the distal half, papilla each (vs. 2-3 papillae in each row, see Fig. 2 and the central light stripe is better developed and B and 26A). In T. caudomaculatum, the second spine reaches almost to the distal margin. The iris is mot- of the first dorsal fin extends much further posteri- tled with black, red and orange pigment with a very orly, reaching to the caudal peduncle. In contrast, clear light stripe margining the dorsal edge of the the second spine of T. tevegae is not elongate, reach- pupil, above which the iris is dark red. A 22.0 mm ing posteriorly only as far as the spine of the second SL female (Fig. 29) is much yellower overall, with dorsal fin when the fin is adpressed. An analysis of the intense yellow pigment in the caudal fin extend- the CO1 gene of 72 specimens originally identified ing almost to the distal margin of the fin and a sprin- as T. tevegae from three localities (Raja Ampat, Palau kling of melanophores in the central light stripe. and Japan) suggests that there are seven deeply diver- The caudal blotch is somewhat better defined, and gent haplogroup clusters, separated from each other disparity in width between the upper and lower by between 9.1-22.9% of the standard mitochondr- halves more obvious. The iris is a mix of golden and ial 5’ COI barcode locus (Winterbottom et al., in black pigment, and the stripe over the pupil is black. prep.). One of these groups, informally called A 23.2 mm SL male (Fig. 30) has an overall reddish “Group 2”, is the species described here as T. xan- cast, grading to orange-red below the dark wedge- thochrum, and it is separated from all the others by shaped internal stripe, and the yellow on the caudal 13.8-17.8%. It is impossible to say at present fin is confined to the proximal half of the fin. whether the true T. tevegae (type locality Rabaul Among the remaining photographed specimens, Harbour, New Britain) is represented in the samples essentially the same variation is apparent, although analysed, since no specimens from that locality are the light stripe under the eye may have a bluish cast. currently available for genetic analysis. The collec- A specimen photographed live underwater at Triton tions analysed to date from both Palau and from Bay (Fig. 31) is overall reddish-orange, with diffuse Raja Ampat contain three deeply divergent hap- body stripes (a somewhat darker mid-lateral stripe logroups, each characterized by extremely low with light orange-yellow dorsal and ventral body within-cluster variation (i. e. < 1%). It is therefore stripes), an oblique white stripe across the iris over possible or even probable that more than one of the top of the pupil, and greyish margins on the these (or other) clusters may be present at the type unbranched rays and a posteriorly attenuating grey locality of T. tevegae, but if so, it is unknown at this stripe in the center of a yellow caudal fin. point which one, if any, might represent the type Colour pattern in alcohol: As above, species. Morphological or colour differences between but pale straw coloured with only the darkly pig- these groups, if they exist, have not been established, mented areas remaining visible. The dark pigmenta- except for T. xanthochrum and the apparent differ- tion on the dorsal surface and the caudal spot are ence in the degree of development of the second dor- especially evident. sal spine between T. caudomaculatum and T. tevegae. Comparisons: This new species is a member of the Distribution: Trimma xanthochrum is currently T. tevegae species group, defined by an interorbital only positively recorded from the Bird’s Head region width of greater than 75% of the pupil width and a of Indonesia (Fig. 6) in depths of 12-70 m. Type A abdominal/caudal vertebral transition. Etymology: The name is derived from the Greek Among the described species in this group, it differs ‘xanthos, yellow or golden, and ‘chros’ colour of the aqua vol. 17 no. 3 - 10 July 2011 160 Richard Winterbottom skin, or body surface, in allusion to the yellow Masuda, H., Amaoka, K., Araga, C., Uyeno, T. & colouration of the body and caudal fin. This species Yoshino, T.). 2nd Edition. Tokai University Press, Tokyo, has been referred to informally (in litt.) as Trimma pp. 236-289. RW sp. 94. AKIHITO, SAKAMOTU, K., IWATA, A., AND IKEDA, Y. 1993. Cephalic sensory organs of the gobioid fishes. In: Fishes of Japan with pictorial keys to the species. (Ed. T. Nakabo). ACKNOWLEDGEMENTS Tokai Univ. Press, Tokyo, Japan [In Japanese], pp 1088- I would especially like to thank Mark V. Erdmann 1116. (Conservation International’s Indonesia Marine AKIHITO, SAKAMOTO, K., IKEDA, Y., & SUGIYAMA, K. 2002. Program) for the invitation to participate in the Gobioidei. In: Fishes of Japan with pictorial keys to the species. fieldwork, and for diligently collecting additional (Ed. T. Nakabo). English edition, Vol. II, Tokai University material after the trip in which I participated. I Press, Tokyo, pp 1139-1310; 1596-1619. would also like to express my gratitude to Laure ALLEN, G. R. & ERDMANN, M. V. 2009. Reef fishes of the Katz who, in addition to collecting many specimens Bird’s Head Peninsula, West Papua, Indonesia. Check List 5 (3): 587-628. of gobies for me, made sure that I always returned ALLEN, G., STEENE, R., HUMANN, P., & DELOACH, N. 2003. safely to the surface after each dive. My sincerest Reef Fish Identification. Tropical Pacific. New World Publi- thanks to the other members of the “fish” team cations Ltd., Florida. 457 pp. (Rudi Dimara, Andi Fauzan, Christine Huffard, AOYAGI, H. 1949. Studies on the fishes of the Riu- Muhammad Lazuardi and Defy Pada), to the cap- Kiu Islands. V. The Zooological Magazine, The Zoological tain and crew of the Putri Papua for their unstinting Society of Japan (Tokyo) 58 (9): 171-173. help, support and companionship during the survey, ASAOKA, R., NAKAE, M. & SASAKI, K. 2011. Description and to the Indonesian Department of Nature Conserva- innervations of the lateral line system in two gobioids, Odontobutis obscura and Pterogobius elapoides (Teleostei: tion (PHKA), the Indonesian Institute of Sciences Perciformes). Ichthyological Research 58: 51-61. (LIPI), and to the Raja Ampat government for their AURICH, H.J. 1938. Mitteilung XXVIII der Wallacea-Expe- support of this work. I also extend my gratitude to dition Woltereck. Die Gobiiden. (Ordnung : Gobioidea). the Paine Trust for their financial support of Internationale Revue der gesamten Hydrobiologie und Hydro- this survey through Conservation International’s graphie, 38(1/2) :125-183. Indonesia Marine Program. Thanks to Doug Hoese COHEN, D. M. & DAVIS, W. P. 1969. Vertical orientation in and Peter Miller for their comments on the section a new gobioid fish from New Britain. Pacific Science 23 (3): on papillae in Trimma, and to Gerry R. Allen and 317-324. Mark V. Erdmann for permission to use their excel- DIMARA, R., FAUZAN, A., LAZUARDI, M., PADA, D., ALLEN, G. R., ERDMANN, M. V., HUFFARD, C. L., KATZ, L. S. & lent photographs. It is always a pleasure to acknowl- WINTERBOTTOM, R. 2010. An illustrated list of additions edge the tremendous contributions of Margaret Zur to the goby fauna of the Raja Ampat Islands, Indonesia (ROM), who for many years gathered and analysed (Pisces, Gobiidae). Check List 6 (4): 619-625. data from the huge collections of borrowed and GOREN, M. 1978. A new gobiid genus and seven new species ROM Trimma. Her notes have been invaluable in from Sinai coasts. Senckenbergiana biologica 59 (3/4): making the comparisons of the new species with 191-203. previously described members of the genus. Addi- GOREN, M. 1981. Three new species and three new records tional financial support for this study was generously of gobies from New Caledonia. Cybium, 3e ser. 5 (3): provided the ROM’s Department of Natural His- 93-101. GOREN, M. 1982. Quisquilius flavicaudatus, a new gobioid tory and my NSERC Discovery Grant A7619. fish from the coral reefs of the Red Sea. Zoologische Med- edelingen 56 (11): 139-142. REFERENCES HAGIWARA, K. & WINTERBOTTOM, R. 2007. Two new AHNELT, H. & BOHACEK, V. 2004. The lateral system of two species of Trimma (Gobiidae) from the western Pacific. Bul- sympatric eastern Pacific gobiid fishes of the genus Lythryp- letin of National Science Museum, Ser. A, Supplement 1: nus (Teleostei: Gobiidae). Bulletin of Marine Science 74 (4): 163-174. 31-51. HAYASHI, M. & SHIRATORI, T. 2003. Gobies of Japanese AHNELT, H. & GÖSCHEL, J. 2003. Morphological differences waters. Hankyu Books, Osaka, Japan (in Japanese). between the eastern Pacific gobiid fishes Quietula guay- HERRE, A. W. C. T. 1945. Notes on fishes in the Zoological masiae and Quietula y-cauda (Teleostei: Gobiidae) with Museum of Stanford University. XIX. — Two new Philip- emphasis on the topography of the lateral-line system. pine gobies, with key to the genera of gobies with vomerine Cybium 27 (3): 185-197. teeth. Proceedings of the Biological Society of Washington AKIHITO, P., HAYASHI , M. & YOSHINO, T. 1988. Suborder 58: 77-82. Gobioidei. In: The Fishes of the Japanese Archipelago. (Eds. HOESE, D. F. 1983. Sensory papillae patterns of the cheek

161 aqua vol. 17 no. 3 - 10 July 2011 Six n. sp. of the genus Trimma (Percomorpha; Gobiidae) from the Raja Ampat Islands, Indonesia, with notes on cephalic sensory papillae nomenclature

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