taxonID	type	description	language	source
039A87C6C062FFE4FF3AFDA4FB14F964.taxon	materials_examined	Material Examined. All type specimens are from the northern part of West Papua province, Indonesia. The description is based on up to 17 specimens, 14.0 – 18.8 mm SL culled from all five lots. Holotype. ROM 94889, (18.4 mm SL female), Mapia Atoll, south end of Pulau Pegun, (00 ° 47.795 ' N, 134 ° 18.198 ' E), steep wall covered with coarse white Halimeda sand, 60 m, clove oil, 2 Nov., 2013, M. V. Erdmann. Paratypes. MZB 22713, 7 (13.5 – 14.7, ex-ROM 87934,), Pulau Mof (north of Waigeo), (00 ° 31.431 ' N, 130 ° 44.020 ' E), 70 m, clove oil, 7 Mar., 2011, M. V. Erdmann. ROM 95269, 5 (14.4 – 16.6), NW side of Ayau Besar Atoll, (00 ° 35.729 ' N, 131 ° 06.212 ' E), exposed oceanic atoll with clear water, steep outer wall with small crevices and overhangs, coarse white coralline sand, 38 m, clove oil, 1 July, 2014, M. V. Erdmann. ROM 98809, 10 (11.0 – 16.8), Ayau Besar Atoll, (00 ° 36.306 ' N, 131 ° 06.398 ' E), steep outer reef slope with white calcareous sand, 40 m, clove oil, 22 Oct., 2014, M. V. Erdmann. ROM 98853, 3 (14.1 – 18.8), NW side of Ayau Besar Atoll, (00 ° 35.177 ' N, 131 ° 06.834 ' E,), 50 m, 19 Feb., 2015, MVE- 15 - 010, M. V. Erdmann. Tissue specimens (all from Ayau Besar Atoll, collected by M. V. Erdmann): MB 0614201, 1 Jul., 2014; MB 0615401, MB 0615402, 22 Oct., 2014; ROM T 19515, 19 Feb., 2015.	en	Winterbottom, Richard, Erdmann, Mark V., Dita Cahyani, N. K. (2015): New species of Trimma (Actinopterygii, Gobiidae) from Indonesia, with comments on head papillae nomenclature. Zootaxa 3973 (2): 201-226, DOI: 10.11646/zootaxa.3973.2.1
039A87C6C062FFE4FF3AFDA4FB14F964.taxon	diagnosis	Diagnosis. A species of Trimma unique in the possession of two dark oblique stripes on either side of the pupil (blue, edged with red in life, dark brown in preservative), the lower of which continues posteriorly across the dorsal margin of the opercle. The dorsal half of the body is red and the ventral half abruptly white in life. The species has a narrow bony interorbital (≤ 50 % pupil width), a moderate interorbital trench with a slight groove posterodorsal to the eye, no scales on the cheek, opercle or in the predorsal midline, no elongate spines in the first dorsal fin, 9 – 10 dorsal and 9 anal fin rays, pectoral fin rays 17 – 18 of which 7 – 11 are branched, and an unbranched fifth pelvic fin ray.	en	Winterbottom, Richard, Erdmann, Mark V., Dita Cahyani, N. K. (2015): New species of Trimma (Actinopterygii, Gobiidae) from Indonesia, with comments on head papillae nomenclature. Zootaxa 3973 (2): 201-226, DOI: 10.11646/zootaxa.3973.2.1
039A87C6C062FFE4FF3AFDA4FB14F964.taxon	description	Description. Dorsal fins VI + I 9 – 10 (mean = 9.6), second or third spine longest but not elongate, reaching posteriorly to between bases of sixth spine of D 1 and spine of D 2 when adpressed, first ray of second dorsal fin unbranched (branched in 3 specimens), posterior element of last dorsal ray unbranched, last ray reaching posteriorly 50 – 67 – 72 % (mean = 65.8 %) of distance between its base and first dorsal procurrent caudal ray; anal fin I 9 (once 10), first ray unbranched as is posterior element of last ray, which reaches posteriorly 45 – 62 – 65 % (mean = 57.4 %) distance between its base and first ventral procurrent caudal ray; pectoral fin 17 – 18 (mean = 17.2), 3 – 4 – 6 (mean = 4.1) dorsal and 2 – 5 (mean = 4.1) ventral unbranched rays separated by 7 – 8 – 12 (mean = 9.1) branched rays, fin reaching posteriorly to a point above area between anus to urogenital papilla; pelvic fin I 5, first four rays with one sequential branch, fifth ray unbranched (Fig. 3) and 52 – 67 % the length of fourth (mean = 57.0, fourth ray damaged in holotype), fourth ray reaching posteriorly to a point between posterior margin of urogenital papilla and base of second anal ray; no fraenum; basal membrane varies from absent, where membrane from fifth ray joins to abdomen lateral to midline to 9 % length of fourth ray (Fig. 3, red arrow). Lateral scales 23 – 24 (mean = 23.4), where 24 count due to additional small anteriormost scale; anterior transverse scales 8 – 9 – 10 (mean = 8.9; yellow arrow in Fig. 3 shows direction of count); posterior transverse scales 7 – 8 (mean = 7.9; blue arrow in Fig. 3 shows direction of count); no scales on median predorsal, cheek or opercle; scales extending anteriorly on sides of body from upper pectoral base curving up to middle of first dorsal fin, a few specimens with 1 – 2 small isolated cycloid scales on side of nape; pectoral-fin base usually with 3 vertical rows of scales, with 1 – 2, 2 – 4 and 3 – 5 scales per row respectively; scales in midline anterior to pelvic fin 4 – 5 (mean = 4.5), where 4, usually an additional scale anteriorly just lateral to midline; scales on pectoral-fin base, breast, anterior midline of belly cycloid (Fig. 3, green arrows show anteriormost ctenoid scales), scales on sides of nape and rest of body ctenoid; 12 circumpeduncular scales, 7 – 8 (mean = 7.2) scales between base of last anal fin ray and first ventral procurrent caudal ray. Upper jaw teeth with an outer row of 6 – 8 evenly spaced, curved, enlarged canines decreasing in size posteriorly and extending to bend of premaxilla, short inner row of straight conical teeth slightly larger than teeth anterior to them, ending before bend of premaxilla, 2 – 3 irregular rows of small conical teeth between these, grading to single row on posterior part of premaxilla. Lower jaw teeth with outer row of about 6 evenly spaced, curved, enlarged canines ending at bend of dentary, inner row of enlarged, slightly curved teeth about one-third length of outer row, decreasing in size posteriorly, ending at beginning of coronoid process, 2 – 3 irregular rows of small conical teeth between these, grading posteriorly to single row and ending near anterodorsal tip of coronoid process of dentary. Tongue broadly rounded, sometimes with small central tip. Gill opening extending anteroventrally to below mid- to posterior third of pupil; outer gill rakers on first gill arch 3 – 4 (mean = 3.3) + 12 – 13 – 14 (mean = 13.1), total rakers 15 – 17 (mean = 16.4). Anterior naris a short tube extending half way over upper lip, posterior naris a pore with raised rim, separated from eye by 1.5 – 2 times its diameter, nasal apparatus occupying anterior two-thirds of snout (Fig. 4). Bony interorbital width 30 – 33 – 43 % (mean = 36.6) of pupil diameter in width, interorbital with moderately developed ‘ U’ - shaped trough about as wide as deep, a shallow postorbital groove but not trench-like. Cephalic sensory papillae as in Figure 4, number of papillae in each row given in Table 1. Ridge of tissue (or dermal crest) may extend anterior to first dorsal spine for up to two-thirds distance to interorbital. Epaxialis extending anteriorly to above a vertical with posterior margin of pupil. Abdominal / caudal vertebral transition not examined, but presumed to be Type B. Colour pattern, live, based on images of holotype, 18.2 mm SL female, ROM 94889, Mapia Atoll, Fig. 5 A, and paratype, Ayau Besar Atoll, ROM 98809, Fig. 5 B. Holotype bicoloured, dark red dorsally and white ventrally below midlateral line continuing anteriorly across cheek just below eye. Midline of dorsum with diffuse white stripe from above posterior margin of pupil to anteriormost dorsal procurrent caudal fin ray. Iris dark gold above pupil, red margined purple stripe across top of pupil which continues anteriorly on snout as diffuse, and much wider, red stripe. White stripe margined dorsally with red across bottom of pupil, continuing anteriorly to maxilla and, after short disjunction behind eye, posteriorly across top of opercle, where white grades to blue margined dorsally with red. Area of snout between stripes and above dorsal stripe yellow, anterior snout and lips red. Rays of dorsal, pectoral and upper half of caudal fins light red (pelvic and anal fin not visible), ventral half of caudal lighter. Upper pectoral fin base with white spot more intense than surroundings. Internal lighter stripe (spinal cord?) anteriorly, followed by three internal lighter spots beneath dorsal fins. Specimen from Ayau Besar Atoll essentially similar, but upper body more orange than red, lower half of body very pale pink, anterior half of lower eye stripe blue rather than white, and spot at upper pectoral fin base more obvious and bluish-white. In other live specimens, dorsum reddish-orange (possibly juveniles). Freshly collected specimen from Pulau Mof (ROM 87934, Fig. 5 C) with yellow head grading to orange on nape and then to orange-red behind this with light greenish-yellow cheek, pale pinkish white abdomen and posterior body, blue or white of iris stripes dark purple, dorsal fins with basal, half-pupil diameter wide basal white stripe bordered dorsally by yellow stripe half this width, then hyaline, caudal fin with outer edges whitish and central rays dirty orange-brown. Preserved colour. Background pale straw yellow, with heavy concentrations of dark brown chromatophores on dorsal half of nape and body, decreasing in number ventrally, where absent from about two scale rows from ventral midline. Moderate number of similar chromatophores on cheek with some on dorsal part of opercle, and on upper and lower lips. Stripes across iris evident as lines of dark brown chromatophores (Fig. 6) which continue anterodorsally onto snout. Lower line continues posterodorsally as dark ovoid patch over dorsal tip of preopercle and from anterodorsal process of opercle half way to its posterodorsal tip; in some specimens dark line continuous from snout to opercle. Fins hyaline except for scattered brown pigment cells on distal half of second dorsal fin, some in caudal fin, and along distal margin of last few anal fin rays.	en	Winterbottom, Richard, Erdmann, Mark V., Dita Cahyani, N. K. (2015): New species of Trimma (Actinopterygii, Gobiidae) from Indonesia, with comments on head papillae nomenclature. Zootaxa 3973 (2): 201-226, DOI: 10.11646/zootaxa.3973.2.1
039A87C6C062FFE4FF3AFDA4FB14F964.taxon	etymology	Etymology. Named in honor of Abraham Aturiri, Governor of West Papua province from 2006 - 2016, in recognition of his energetic efforts to ensure the conservation and wise, sustainable use of the hugely diverse coral reefs of the Bird's Head Seascape for the benefit of the Papuan people. The species has been informally referred to as Trimma RW sp 99.	en	Winterbottom, Richard, Erdmann, Mark V., Dita Cahyani, N. K. (2015): New species of Trimma (Actinopterygii, Gobiidae) from Indonesia, with comments on head papillae nomenclature. Zootaxa 3973 (2): 201-226, DOI: 10.11646/zootaxa.3973.2.1
039A87C6C062FFE4FF3AFDA4FB14F964.taxon	distribution	Distribution. Currently known with certainty only from the northern oceanic atolls of West Papua province, Indonesia (Fig. 18). Comparisons. Trimma aturirii is unique among species in the genus in its bipartite colour pattern (red or orange above, white or pale pink below). The two oblique dark stripes on either side of the pupil (blue-edged with red margins in life, dark brown in preservative), the lower of which continues after a short hiatus posteriorly along the dorsal margin of the opercle, are shared only with an undescribed species (T. RW sp 49) from the Line Islands, which lie some 7,500 kilometers due east of the easternmost sample from Indonesia. Specimens from the latter locality invariably have 10 (vs. 9) anal rays, and the red colouration of the body extends ventrally almost to the ventral midline (vs. confined above midlateral septum). Ten other described species of Trimma share with the new species the lack of scales in the predorsal midline and an unbranched fifth pelvic fin ray. Of these, T. anaima Winterbottom, 2000, has 8 dorsal and anal fin rays, unbranched pectoral fin rays, and less than 15 total gill rakers (vs. 9 – 10, 9, middle rays branched and 15 or more gill rakers). Trimma grammistes (Tomiyama, 1936) has unbranched pectoral fin rays and a second dorsal spine that reaches posteriorly beyond the first element of the second dorsal fin when adpressed (vs. branched, and not reaching beyond spine). That species also has a pupildiameter wide black stripe from the upper lip through the eye and along the body above the midlateral line until it curves gradually to the center of the caudal peduncle and onto the caudal fin (vs. stripe absent). Three other species, T. bisella Winterbottom, 2000, T. cana Winterbottom, 2004, and T. sostra Winterbottom, 2004, have greatly elongate second dorsal spines (vs. not elongate) and lack a dermal crest anterior to the first dorsal fin (vs. present). The former species has four red bars on the head and white saddles on the peduncle (vs. both absent), while the latter two have a pale body with seven broad red bars or blotches (vs. bars absent). Trimma sanguinellus Winterbottom & Southcott, 2007 has an entirely red body, unbranched pectoral fin rays, and a second dorsal spine that extends posteriorly beyond the first element of the second dorsal fin when adpressed (vs. bicoloured, branched, and second spine anterior first spine of second dorsal fin respectively). Trimma maiandros Hoese et al., 2011, has a pale grey body with large yellow blotches, 5 or fewer branched pectoral rays, and no opercular stripe (vs. bicoloured, <5 branched pectoral rays, and dark opercular stripe respectively). Trimma stobbsi Winterbottom, 2001 has a yellow to brown body with scale pockets outlined and a dark, half pupil diameter sized, roughly ovoid spot on the upper margin of the opercle near its posterior margin (vs. bicoloured with stripe along dorsal margin of opercle). Trimma hotsarihiensis Winterbottom, 2009, has a yellow body with scale margins more strongly marked, yellow spots in the caudal fin and dark basal stripes in the dorsal and anal fins. It has a dark diffuse stripe across the top of the opercle, but the stripe is horizontal and ends anteriorly at the level of the mid-pupil rather than being obliquely oriented downwards to pass below the eye. Trimma kudoi Suzuki & Senou, 2008, has a similar overall colouration as T. hotsarihiensis, but lacks the dark stripe over the opercle. It has two oblique purple-blue stripes, one at the dorsal margin of the iris and a second that passes across the middle of the pupil (vs. two stripes on either side of the pupil).	en	Winterbottom, Richard, Erdmann, Mark V., Dita Cahyani, N. K. (2015): New species of Trimma (Actinopterygii, Gobiidae) from Indonesia, with comments on head papillae nomenclature. Zootaxa 3973 (2): 201-226, DOI: 10.11646/zootaxa.3973.2.1
039A87C6C062FFE4FF3AFDA4FB14F964.taxon	discussion	Discussion. Specimens of this species were usually found perched individually on vertical surfaces at the back of crevices and overhangs in depths ranging between 38 – 70 m. Despite extensive sampling of these depths on reefs throughout eastern Indonesia, this species has only been found on exposed outer reef slopes of the northernmost oceanic atolls and islands of West Papua (Mapia Atoll, Ayau Besar Atoll and Pulau Mof). An analysis of the CO 1 gene of the three tissue specimens of this species suggests that they are phenetically closest to T. hotsarihiensis, from which they are separated by a minimum of 9.9 % of the base pairs.	en	Winterbottom, Richard, Erdmann, Mark V., Dita Cahyani, N. K. (2015): New species of Trimma (Actinopterygii, Gobiidae) from Indonesia, with comments on head papillae nomenclature. Zootaxa 3973 (2): 201-226, DOI: 10.11646/zootaxa.3973.2.1
039A87C6C069FFFDFF3AFF5CFE3CFF7C.taxon	materials_examined	Material examined. Five lots, five specimens (12.0 – 16.7 mm SL), description excludes data from ROM T 012726 (tissue specimen), all from the West Papua province of Indonesia. Holotype. ROM 98815, (16.7), Kaimana, Triton Bay, White Rock Falls, (03 ° 53.757 ' S, 134 ° 06.638 ' E), 50 m, clove oil, field number: MVE- 14 - 018, 3 Dec., 2014, M. V. Erdmann. Paratypes. MZB 22714, (12.0, ex-ROM 936491), Teluk Cendrawasih, Pulau Mansinam, (00 ° 55.344 ' S, 134 ° 06.557 ' E), 70 m, clove oil, 20 Mar., 2012, M. V. Erdmann. ROM 87545, (15.8), Raja Ampat, Keruo I., (00 ° 35.269 ' S, 130 ° 17.685 ' E), steep slope with overhangs sand / gorgonians, 70 m, clove oil, 27 Aug., 2010, M. V. Erdmann. ROM T 012726, (12.2), Raja Ampat, Keruo I., (00 ° 35.269 ' S, 130 ° 17.685 ' E), steep slope with overhangs sand / gorgonians, 70 m, clove oil, 6 Dec., 2010, M. V. Erdmann. ROM 88107, (13.8), Raja Ampat, off Misool I., Pulau Fiabacet, (02 ° 13.196 ' S, 130 ° 33.834 ' E), clove oil, 26 Oct., 2010, M. V. Erdmann.	en	Winterbottom, Richard, Erdmann, Mark V., Dita Cahyani, N. K. (2015): New species of Trimma (Actinopterygii, Gobiidae) from Indonesia, with comments on head papillae nomenclature. Zootaxa 3973 (2): 201-226, DOI: 10.11646/zootaxa.3973.2.1
039A87C6C069FFFDFF3AFF5CFE3CFF7C.taxon	diagnosis	Diagnosis. A species of Trimma with a narrow bony interorbital (≤ 40 % pupil width), a moderate interorbital trench with a slight groove posterodorsal to the eye, 17 – 18 unbranched pectoral fin rays, 1 – 5 cycloid scales in the predorsal midline confined to about the middle third of the distance between the posterodorsal margin of the eye and the base of the spine of the first dorsal fin, and a single row of 1 – 3 cycloid scales along the upper border of the opercle. In live and freshly collected specimens of this species, the presence of oval red hyoid spots connected anteriorly in the midline is diagnostic.	en	Winterbottom, Richard, Erdmann, Mark V., Dita Cahyani, N. K. (2015): New species of Trimma (Actinopterygii, Gobiidae) from Indonesia, with comments on head papillae nomenclature. Zootaxa 3973 (2): 201-226, DOI: 10.11646/zootaxa.3973.2.1
039A87C6C069FFFDFF3AFF5CFE3CFF7C.taxon	description	Description. Based on the holotype and an additional three specimens (12.0 – 16.7 mm SL). Dorsal fins VI + I 9, second or third spine longest but not elongate, and reaching to between base of first to second elements of D 2 when adpressed, first ray of second dorsal fin branched (unbranched in holotype), posterior element of last dorsal ray unbranched, last ray reaching posteriorly 55 – 64 % (mean = 59.0 %) of distance between its base and first dorsal procurrent caudal ray; anal fin I 8 – 9, first ray unbranched, as is posterior element of last ray, which reaches posteriorly 40 – 48 – 54 % (mean = 47.3 %) distance between its base and first ventral procurrent caudal ray; pectoral fin 17 – 18 (mean = 17.8), all rays unbranched, fin reaching posteriorly to a point above first to third anal fin element; pelvic fin I 5, first four rays with one sequential branch, fifth ray unbranched (Fig. 7 A) or branched (n = 1, Fig. 7 B) and 52 – 60 % the length of fourth (mean = 56.5 %), fourth ray reaching posteriorly to a point between bases of third to sixth anal elements; no fraenum; basal membrane varies from vestigial (perhaps torn) to 11 – 15 % length of fourth ray (Fig. 7 B). Lateral scales 23; anterior transverse scales 8 – 9 (mean = 8.5); posterior transverse scales 7 – 8 (mean = 7.3); median predorsal scales 1 – 5 (mean = 2.8; Fig. 8); scaled midline area confined to about middle third of nape with naked areas anteriorly and posteriorly; scales extending anteriorly on sides of nape to just behind eye; scales on cheek apparently absent; opercle with single row of 1 – 3 cycloid scales; pectoral-fin base margined posteriorly with 2 – 3 vertical rows of cycloid scales with 4 scales in outer row, 3 in row anterior to this, and 1 in anterior-most row (if present, holotype only); scales anterior to pelvic fin 4 – 6 (mean = 4.8), scales on and immediately behind pectoral-fin base, breast, anterior midline of belly, and midline of nape cycloid, scales on sides of nape may be ctenoid or cycloid, other scales ctenoid. Upper jaw teeth with an outer row of about 6 evenly spaced, curved, enlarged canines decreasing in size posteriorly and extending to bend of premaxilla, inner row of about 4 straight conical teeth about one-third size of outer row, ending before bend of premaxilla, 2 – 3 irregular rows of small conical teeth between these, grading to single row on posterior part of premaxilla. Lower jaw teeth with outer row of about 6 evenly spaced, curved, enlarged canines ending at bend of dentary, inner row of slightly enlarged and curved teeth decreasing in size posteriorly to end at beginning of coronoid process, 2 – 3 irregular rows of small conical teeth between these, grading posteriorly to single row ending near anterodorsal tip of coronoid process of dentary. Tongue teretely rounded to weakly bilobed. Gill opening extending anteroventrally to below mid- to posterior third-pupil; outer gill rakers on first gill arch 3 – 4 (mean = 3.3) + 13 – 14 (mean = 13.5), total rakers 16 – 17 (mean = 16.7). Anterior naris a long thin tube extending well past upper lip, almost equal in length to width of nasal capsule, which is confined to anterior half of snout; posterior naris a pore with raised rim, separated from eye by 2.5 – 3 times its diameter (Fig. 9). Bony interorbital width 34 – 40 % of pupil diameter in width (mean = 38.0 %), interorbital with moderately developed ‘ U’ - shaped trench about as wide as deep, a shallow postorbital groove but not trench-like. Sensory papillae as in Figure 9, number of papillae in each row given in Table 1. Ridge of tissue (or dermal crest) may extend anterior to first dorsal spine for one-third distance to interorbital (Fig. 8 A, red arrows). Epaxialis extending anteriorly to above a vertical with posterior margin of pupil. Abdominal / caudal vertebral transition not examined, but presumed to be Type B. Colour pattern, freshly collected, based on images of holotype, 16.7 mm SL male, ROM 98815, Fig. 10 A, and paratype, 12.2 mm SL, ROM T 012726, Fig. 10 B. Holotype with head light purplish-grey, grading into light yellow behind eye, along dorsal margin of opercle and across posterior margin of opercle (where suffused with pale red), light pink lips, upper branchiostegal membrane and pectoral fin base, cheek with scattered iridocytes and darker pigment cells; ovoid bright red spot with diffuse dark pigment cells centrally over hyoid arch on ventral surface, tapering anteriorly where joined at midline below hypohyals, diverging slightly from midline posteriorly (Fig. 11 A). Iris with dark blue border and thin, sandy-yellowish halo around pupil, middle portion reddish with darker diffuse stripe above pupil. Nape and body greenish yellow dorsally, becoming less green below midlateral septum, posterior margins of scales slightly darker, especially on anterodorsal half of body, some scattered dark chromatophores present, especially anteriorly. First dorsal fin with half-pupil diameter basal yellow stripe, fin membrane above this hyaline with numerous scattered iridocytes. Second dorsal fin with narrow dark basal stripe followed by yellow stripe (in line with stripe of first dorsal fin), membrane above this with iridocytes, fin rays pinkish. Anal fin mostly yellow, margined by distal stripe of iridocytes and dark pigment cells. Caudal fin yellow, somewhat paler distally. Pectoral fin rays yellow, membrane hyaline. Pelvic fin rays off-white, scattered iridocytes in membrane. Paratype similar (Fig. 10 B), but snout, jaws and anterior cheek red, middle ring of iris mix of red and yellow, no darker stripe above pupil, scale pockets on side of nape outlined with darker pigment, breast and lower pectoral fin base orange and numerous iridocytes on belly. Preserved colour. Holotype pale straw-yellow, cheek and opercle with heavy scattering of dark chromatophores, sides and top of snout covered with small round chromatophores with some scattered on lips, nape with scattered melanophores and larger brown chromatophores, which decrease in concentration along dorsum to end of second dorsal fin, and tend to be concentrated along posterior margins of scales. Diffuse hyoid spot made up of brown chromatophores on branchiostegal membrane beneath hypohyals and below posterior half of eye (Fig. 11 B). Base and distal half of first dorsal fin hyaline, with stripe of dark chromatophores between; thin dark basal stripe in second dorsal fin followed by clear stripe and then another diffuse dark stripe before becoming hyaline distally; anal fin with similar basal and distal dark stripes, middle of fin hyaline; caudal fin hyaline with few scattered dark chromatophores (especially on ventral procurrent fin rays); pectoral fin hyaline; pelvic fin with some dark chromatophores along margins of rays. Paratype similar, but with dark chromatophores on nape and anterior body more strongly associated with scale pockets, and no trace of hyoid spot.	en	Winterbottom, Richard, Erdmann, Mark V., Dita Cahyani, N. K. (2015): New species of Trimma (Actinopterygii, Gobiidae) from Indonesia, with comments on head papillae nomenclature. Zootaxa 3973 (2): 201-226, DOI: 10.11646/zootaxa.3973.2.1
039A87C6C069FFFDFF3AFF5CFE3CFF7C.taxon	etymology	Etymology. Named for the Greek ‘ kardia’, a heart, in allusion to the two ovoid red spots that are joined anteriorly on the ventral surface of the head, and have an admittedly tenuous stereotypical resemblance to that organ when viewed from below. This species has been informally referred to as Trimma RW sp 98.	en	Winterbottom, Richard, Erdmann, Mark V., Dita Cahyani, N. K. (2015): New species of Trimma (Actinopterygii, Gobiidae) from Indonesia, with comments on head papillae nomenclature. Zootaxa 3973 (2): 201-226, DOI: 10.11646/zootaxa.3973.2.1
039A87C6C069FFFDFF3AFF5CFE3CFF7C.taxon	distribution	Distribution. Currently know from 5 specimens collected in 50 – 70 m at Raja Ampat, Cendrawasih Bay and Triton Bay, all in the province of West Papua, Indonesia (Fig. 18). Comparisons. Most species of Trimma either have a scaled or a naked predorsal midline. However, several species exhibit variation in this region. A few (<6 %) specimens of T. maiandros Hoese et al., 2011, may have 1 – 5 scales in the middle of the nape as in the present species. That species lacks opercular scales (vs. present), has a dark zig-zag stripe on the nape and dorsal half of the body separated by yellow blotches, which are also present on the nape and cheek (vs. such markings absent), and lacks the red hyoid spots (vs. present). Some specimens of Trimma anthrenum Winterbottom, 2006, may have reduced predorsal scalation with naked areas in front of the first dorsal spine and immediately behind the eye. Trimma anthrenum has at least some branched pectoral fin rays (vs. unbranched), and lacks scales on the opercle and the red hyoid spot (vs. both present). A few large specimens (> 20 mm SL) of T. erdmanni Winterbottom, 2011, have up to 6 scales in the midline of the central portion of the predorsal area, but may be differentiated by the presence of branched pectoral fin rays, a red lateral stripe on the body (vs. both absent) and by the lack of the red hyoid spots. Trimma hoesei Winterbottom, 1984, has a few scales in the predorsal midline, but these are confined to the area just anterior to the first dorsal fin spine (vs. middle of the nape), there are branched pectoral fin rays (vs. unbranched), no opercular scales (vs. present) and the caudal fin is forked (vs. rounded). Trimma okinawae (Aoyagi, 1949) and T. readerae Winterbottom & Hoese, 2015 may have 3 – 5 scales across the midline of the nape anterior to the first dorsal fin spine, and additionally may have a red spot on the branchiostegal membranes. However, these two species have branched rays in the pectoral fin (vs. unbranched), lack opercular scales (vs. present) and the red spot on the branchiostegal membranes are smaller, situated more posteriorly (just below the vertical limb of the preopercle and posterior to the point where the branchiostegal membranes join in the midline), and do not join anteriorly in the midline. Trimma irinae Winterbottom, 2014 also has a dark red spot on the branchiostegal membranes, but has a fully scaled predorsal midline (vs. partially scaled) an elongate second dorsal spine (vs. not elongate), branched pectoral fin rays (vs. unbranched) and two black spots near the base of the first dorsal fin (vs. absent). Trimma hayashii Hagiwara & Winterbottom, 2007 has a dark ocellated spot on the branchiostegal membranes, but its position is the same as that in T. okinawae and T. readerae above (vs. anterior to the junction of the membranes). It also has some pectoral fin rays branched (vs. unbranched), lacks opercular scales (vs. present) and has red spots on the head and nape (vs. spots absent).	en	Winterbottom, Richard, Erdmann, Mark V., Dita Cahyani, N. K. (2015): New species of Trimma (Actinopterygii, Gobiidae) from Indonesia, with comments on head papillae nomenclature. Zootaxa 3973 (2): 201-226, DOI: 10.11646/zootaxa.3973.2.1
039A87C6C069FFFDFF3AFF5CFE3CFF7C.taxon	discussion	Discussion. Specimens examined for this study were collected between 50 – 71 m, where they were invariably collected living associated with larger coral rubble pieces lying on soft bottoms below the reef slope. Unlike most species of Trimma, this species appears to live a solitary existence, and only single specimens were collected at any given time. The species seems to have a similar behaviour to T. meranyx Winterbottom et al., 2014 a, which cooccurs with this species and has a similar depth distribution. Other deep-dwelling Trimma that also are generally found individually include T. zurae Winterbottom et al., 2014 a and T. irinae. Phenetically, the single specimen of this species available for genetic analysis forms a grade with an undescribed species from Fiji (T. RW sp 97) and three haplogroups of T. stobbsi according to the CO 1 analysis of Winterbottom et al. (2014 b).	en	Winterbottom, Richard, Erdmann, Mark V., Dita Cahyani, N. K. (2015): New species of Trimma (Actinopterygii, Gobiidae) from Indonesia, with comments on head papillae nomenclature. Zootaxa 3973 (2): 201-226, DOI: 10.11646/zootaxa.3973.2.1
039A87C6C070FFF5FF3AF94BFE70FEE1.taxon	materials_examined	Material. The type material is restricted to specimens collected in Cendrawasih Bay, which forms the eastern and northwestern margins of West Papua and Papua provinces, Indonesia, respectively. Holotype. ROM 98816, 16.5 mm SL female, Indonesia, West Papua, Cendrawasih Bay, Mark's Delite, about 45 kms E of Pulau Roon, (02 ° 25.833 ' S, 134 ° 59.366 ' E), 20 m, clove oil, 13 May, 2014, M. V. Erdmann. Paratypes. MZB 22715, 15 (16.7 – 17.5, ex- WAM P. 33043.001), Tridacna Atoll, outer reef, (02 ° 30 ' S, 134 ° 59 ' E), 20 – 60 m, CB 08 – 005, 8 Nov., 2008, G. R. Allen & M. V. Erdmann. ROM 95249, 13 (11.3 – 19.4), collected with holotype. ROM 93648, 6 (14.9 – 17.2), Mark’s Treasure, (02 ° 25.830 ' S, 134 ° 59.409 ' E), 45 m, clove oil, 15 Jun., 2012, M. V. Erdmann. ROM 95778, 2 (13.1 – 14.7), Tridacna Atoll, (02 ° 29 ' 37.6 " S, 134 ° 58 ' 59.4 " E), 50 m, clove oil, 15 Jul., 2013, M. V. Erdmann. WAM P. 33043.001, 15 (15.4 – 18.2), data as for MZB 22715. WAM P. 33357.007), 13 (9.4 – 18.0), Tridacna Atoll, outer reef, (02 ° 30 ' S, 134 ° 59 ' E), 70 – 71 m, CB 10 - 002, 1 Sept., 2008, G. R. Allen & M. V. Erdmann. Non-type material. Indonesia: Bali: ROM 94022, (17.5), Pulau Menjangan, (08 ° 05.813 ' S 114 ° 31.608 ' E), 8 m, 14 Sept., 2012, M. V. Erdmann. ROM T 012754, (20.2), as for ROM 94022. Sulawesi: ROM 94884, 5 (16.8 – 21.9), North Sulawesi, Bunaken I., Fukui, cave with silt, (01 ° 36.755 ' N, 124 ° 44.333 ' E), 70 m, 2 Sept., 2013, M. V. Erdmann. Palau: ROM 74797, 5 (15.2 – 17.1), Uchelbeluu Reef (aka Short Drop Off), (07 ° 16 ' 29.2 " N, 134 ° 31 ' 32.1 " E), 14 – 26 m, 21 May, 2004, R. Winterbottom et al. ROM 80393, (19.8), Uchelbeluu Reef, (07 ° 16 ' 28.7 " N, 134 ° 31 ' 32 " E), 20 – 27 m, 31 Mar., 2006, R. Winterbottom et al. Papua New Guinea: New Britain, Rabaul: ROM 92317, (16.2), Dawapia Rocks, (04 ° 14.185 ' S, 152 ° 09.980 ' E), 17 m, 26 Nov., 2011, R. Winterbottom & W. Holleman. ROM T 13200, (16.2), as for ROM 92317. WAM P. 28170 – 004, (17.7), six kilometers south east of Rabaul, near Bai village at steep dropoff (04 ° 11 ' S, 152 ° 12 ' E), 25 – 34 m, 0 7 Oct., 1983, G. Allen. Hermit Is, Amot I. USNM 263453, 2 (13.1 – 15.2), ocean side of reef on drop-off (01 ° 33 ' S, 144 ° 59 ' E), 0 – 46 m, 31 Oct., 1978, Springer et al. Philippines: Balacasag I. USNM 258622, (18.9), west side at drop-off (09 ° 31 ' 14 " N, 123 ° 40 ' 00 " E), 0 – 24.4 m, 10 Jun., 1978, Springer et al. USNM 263467, 7 (14.0 – 20.6), W side at drop-off, (09 ° 31 ' 14 " N, 123 ° 40 ' 00 " E), 0 – 24 m, 10 Jun., 1978, Springer et al. USNM 263468, (19.1), W side at drop-off, (09 ° 31 ' 14 " N, 123 ° 40 ' 00 " E), 0 – 41 m, 11 Jun., 1978, Springer et al. Cebu: Caceres Reef: USNM 263521, 5 (15.1 – 18.6), near Huisan Point, east coast of Cebu, 24.4 – 30.5 m, 18 May 1979, J. Libbey et al. Cebu: Mactan I: ROM 49217, (13.2), off Hudson Beach, (10 ° 15 ' N, 124 ° 00 ' E), 12 – 20 m, 7 Aug., 1985, R. Winterbottom & E. O. Murdy. ROM 49218, 2 (8.8 – 17.7), off Hudson Beach, (0 ° 15 ' N, 124 ° 00 ' E), 12 – 18 m, 8 Aug., 1985, R. Winterbottom & E. O. Murdy. Negros Oriental: ROM 53062, (17.5), Bais Bay near main channel, (09 ° 36 ' 54 " N, 123 ° 11 ' 06 " E), 18 – 35 m, 19 May, 1987, R. Winterbottom et al. Siquijor Island: ROM 53059, (20.9), Tonga Point, (09 ° 12 ' 16 " N, 123 ° 26 ' 48 " E), 14 – 18 m, 8 May, 1987, R. Winterbottom et al. ROM 53060, (18.2), Tonga Point, (09 ° 12 ' 16 " N, 123 ° 26 ' 48 " E), 5 – 12 m, 9 May, 1987, G. D. Johnson et al. ROM 53061, 10 (17.2 – 21.8, one specimen 20.3, cleared and stained, ROM 1410 CS), Tonga Point, (09 ° 12 ' 16 " N, 123 ° 26 ' 48 " E), 15 – 21 m, 12 May, 1987, R. Winterbottom et al. ROM 53064, 3 (17.9 – 19.0), Tonga Point, (09 ° 12 ' 16 " N, 123 ° 26 ' 48 " E), 9 – 21 m, R. Winterbottom et al., 22 May, 1987. USNM 263561, 2 (18.0 – 20.0), 2 km W of town of Siquijor, (09 ° 13 ' 30 " N 123 ° 29 ' E), 24.4 – 30.5 m, 14 May 1979, J. Libbey et al.	en	Winterbottom, Richard, Erdmann, Mark V., Dita Cahyani, N. K. (2015): New species of Trimma (Actinopterygii, Gobiidae) from Indonesia, with comments on head papillae nomenclature. Zootaxa 3973 (2): 201-226, DOI: 10.11646/zootaxa.3973.2.1
039A87C6C070FFF5FF3AF94BFE70FEE1.taxon	diagnosis	Diagnosis. A species of Trimma with a large (slightly greater than pupil diameter in width) round, black, ocellated spot in the first dorsal fin between spines 1 and 5, a second, much smaller black or dark red spot just posterior to the spine of the second dorsal fin and above the basal stripe, a yellow body with a dark purplish or gray head with two distinct red bars across the cheek, no round spots of any colour on the nape, opercle or cheek, a small dark (preserved) or white (alive) spot on the upper pectoral fin base, a narrow bony interorbital (<70 % pupil diameter), no elongate spines in the first dorsal fin, 15 – 16 pectoral fin rays with the middle 4 – 8 rays branched, a fifth pelvic fin ray with a single dichotomous branch, and 16 – 17 total gill rakers on the first gill arch.	en	Winterbottom, Richard, Erdmann, Mark V., Dita Cahyani, N. K. (2015): New species of Trimma (Actinopterygii, Gobiidae) from Indonesia, with comments on head papillae nomenclature. Zootaxa 3973 (2): 201-226, DOI: 10.11646/zootaxa.3973.2.1
039A87C6C070FFF5FF3AF94BFE70FEE1.taxon	description	Description. Based on the holotype and 10 specimens (15.4 – 19.4 mm SL) from ROM 95249. Dorsal fins VI + I 8 – 9 (mean = 8.8), second spine longest and reaching to between base of first to third elements of D 2 when adpressed, first ray of second dorsal fin unbranched, (once branched), posterior element of last dorsal ray unbranched, last ray reaching posteriorly 28 – 37.4 – 44 % (mean = 35.7 %) of distance between its base and first dorsal procurrent caudal ray; anal fin I 8, first ray unbranched, as is posterior element of last ray, which reaches posteriorly 29 – 34.8 – 43 % (mean = 33.8 %) distance between its base and first ventral procurrent caudal ray; pectoral fin 15 – 16 (mean = 15.8), 4 – 7 dorsal and 4 – 5 – 6 ventral rays unbranched, total unbranched rays 8 – 9 – 12 (mean = 9.7) with 4 – 7 – 8 (mean = 6.1) branched rays, fin reaching posteriorly to above first, second or third anal fin element; pelvic fin I 5, first four rays with one sequential branch, fifth ray unbranched and 41 – 43 – 54 % length of fourth (mean = 47.7 %), fourth ray reaching posteriorly to between bases of first to sixth anal fin elements; no fraenum; basal membrane vestigial (<7 % length of fourth ray — Fig. 12, red arrow) or not crossing midline and attached to side of body beneath last pre-pelvic scale. Lateral scales 23; anterior transverse scales 8 (once 9); posterior transverse scales 7 – 8; predorsal scales 7 – 8 (mean = 7.7); scales extending anteriorly on sides of nape to just behind eye, scales on cheek apparently absent; opercle with 1 – 2 rows of 3 and 1 – 2 cycloid scales respectively or with larger scales ctenoid; pectoral-fin base margined posteriorly with two vertical rows of 3 – 4 scales in outer row and 1 – 2 in inner row; scales anterior to pelvic fin 6 – 7 (mean = 6.7), scales on pectoral-fin base, breast, anterior midline of belly and anterior nape cycloid, others ctenoid. Upper jaw teeth with outer row of about 8 evenly spaced, curved, enlarged canines decreasing in size posteriorly and extending almost to distal end of premaxilla, inner row of about 4 straight conical teeth about half size of outer row, ending before bend of premaxilla, 2 – 3 irregular rows of small conical teeth between these, grading to single row on posterior part of premaxilla. Lower jaw teeth with outer row of about 6 evenly spaced, curved, enlarged canines ending at bend of dentary, inner row of slightly enlarged and curved teeth which decrease in size posteriorly and end at coronoid process, 2 – 3 irregular rows of small conical teeth between these, grading posteriorly to single row and ending on anterodorsal tip of coronoid process of dentary. (Description of teeth based on a single, non-type cleared and stained specimen from the Philippines, ROM 1410 CS). Tongue broadly truncate, often slightly concave in midline. Gill opening extending anteroventrally to below mid-pupil; outer gill rakers on first gill arch 3 – 4 (mean = 3.2) + 13 – 14 (mean = 13.6), total rakers 16 – 17 – 18 (mean = 16.8). Anterior naris short tube, separated from upper lip by its width, post naris small pore with a slight or no raised rim and more or less flush with surface of snout, separated from eye by 3 – 4 times its diameter, nasal apparatus small, confined to anterior one-third of snout (Fig. 13 B). Bony interorbital width 37 – 40 – 47 % of pupil diameter in width (mean 41.1 %), interorbital with moderately developed ' U' - shaped trench about as wide as deep, a shallow postorbital groove but not trench-like. Sensory papillae as in Figure 13, number of papillae in each row given in Table 1. No ridge of tissue (or dermal crest) extending anterior to first dorsal spine. Epaxialis reaching anteriorly to above vertical with posterior margin of pupil. Abdominal / caudal vertebral transition Type B, based on non-type, cleared and stained specimen (ROM 1410 CS) from the Philippines (Fig. 14). Colour pattern, live and freshly collected, based on numerous photographs by G. R. Allen and M. V. Erdmann (Fig. 15). Head (except opercular region) dark purplish-gray, lighter in juveniles, with two one-third pupil diameter red bars across cheek, first bar below anterior margin of pupil, second bar below posterior margin of pupil and may be slightly convex along its anterior margin; faint diffuse reddish marking may be present between eye and maxilla, remainder of cheek dark grey or purple (lighter in juveniles) sprinkled with dark chromatophores, larger within red bars than elsewhere on cheek; iris mottled with dark red and black or purple with thin white circle around iris; two to three red spots separated by lighter bluish-purple interspaces along dorsomedial margin. Opercular region and body yellow, scales on nape and upper body below first dorsal fin usually edged with darker pigment (less so in juveniles). First dorsal fin with large black spot (slightly greater in diameter than pupil) between spines 1 – 5, margined anteriorly and dorsally with white or light blue; remainder of fin yellowish with a diffuse basal band of iridocytes and dark chromatophores; distal margin of fin between spines 4 – 6 may also have iridocytes and dark chromatophores. Second dorsal fin with similar basal and distal stripes, main body of fin translucent yellow; dark red or black round spot almost fills space between spine and first ray just above distal stripe. Caudal fin translucent, often with yellowish wash. Anal translucent yellow with leading and distal margins dirty white or pale blue. Pelvic and pectoral membranes translucent, pectoral rays yellow to reddish; white spot diffusely margined with melanophores on upper pectoral fin base. Preserved colour. Background colour of head and body off-white (Fig. 16). Cheek with many dark chromatophores except where red bars occurred, where pale with only a few chromatophores. Top of snout very heavily pigmented with dark chromatophores, nape and opercular region somewhat less so, with pigment concentrated on, but not confined to, scale margins on nape; behind this, dark chromatophores along scale margins with a few others scattered on body, amount of pigmentation decreasing posteriorly and ventrally. Large black spot in first dorsal still evident, but rest of fin hyaline except for a distal dusting of dark chromatophores. Black (or sometimes red in life) spot behind spine of second dorsal fin also present, as are dark chromatophores in basal stripe and along distal margin of fin. Caudal fin translucent with a few scattered chromatophores. Anal fin hyaline with darker basal and distal stripes. Pelvic fin with a few darker pigment cells basally. Pectoral fin membranes hyaline, a small concentration of dark chromatophores forming diffuse spot on dorsal base of fin.	en	Winterbottom, Richard, Erdmann, Mark V., Dita Cahyani, N. K. (2015): New species of Trimma (Actinopterygii, Gobiidae) from Indonesia, with comments on head papillae nomenclature. Zootaxa 3973 (2): 201-226, DOI: 10.11646/zootaxa.3973.2.1
039A87C6C070FFF5FF3AF94BFE70FEE1.taxon	etymology	Etymology. The name is derived from “ tri ” (three), and “ oculatus ” (having eyes) in reference to the extra apparent eye formed by the dark ocellated spot in the first dorsal fin. This species has been informally referred to as Trimma RW sp. 32.	en	Winterbottom, Richard, Erdmann, Mark V., Dita Cahyani, N. K. (2015): New species of Trimma (Actinopterygii, Gobiidae) from Indonesia, with comments on head papillae nomenclature. Zootaxa 3973 (2): 201-226, DOI: 10.11646/zootaxa.3973.2.1
039A87C6C070FFF5FF3AF94BFE70FEE1.taxon	distribution	Distribution. Positively known only from the northern portion Cendrawasih Bay, West Papua, Indonesia (see Discussion below, Fig. 18). Comparisons. Only one other species of Trimma, T. zurae has a large, round, ocellated black spot in the first dorsal fin. In T. zurae, the black spot is about equal to the eye diameter in width (vs. slightly greater than pupil diameter), there are yellow spots on the nape, posterodorsal cheek and upper opercle (vs. such spots absent), there are no red bars on the cheek (vs. two such bars present), and there are 17 – 18 pectoral fin rays (vs. 15 – 16). In addition, the posterior nares of T. trioculatum lack a raised rim, and the nasal sac is not elevated above the surrounding surface of the snout (vs. posterior nares with a raised rim, nasal sac slightly raised above surrounding snout in T. zurae). In T. irinae there is an ovoid dark basal spot in the first dorsal fin between spines 1 – 4, with a smaller rounded basal spot between spines 5 – 6. This species lacks the dark spot behind the spine of the second dorsal fin, has an elongate second dorsal spine reaching posteriorly to the end of, or beyond the second dorsal fin (vs. second spine not elongate), has round reddish spots on the posterior head and anterior body (vs. such spots absent), lacks bars on the cheek (vs. present), has a dark spot on the branchiostegal membranes (vs. such a spot absent), and has more total gill rakers (19 – 20 vs. 16 – 18). Trimma papayum Winterbottom, 2011, also has one or two dark spots near the base of the first dorsal fin, but these are very small (about one-quarter pupil diameter in width vs. a single spot greater than pupil diameter in width), an elongate second dorsal spine in the first dorsal fin (vs. not elongate), orange spots on the opercle and nape (vs. such spots absent), and 18 – 19 pectoral fin rays (vs. 15 – 16).	en	Winterbottom, Richard, Erdmann, Mark V., Dita Cahyani, N. K. (2015): New species of Trimma (Actinopterygii, Gobiidae) from Indonesia, with comments on head papillae nomenclature. Zootaxa 3973 (2): 201-226, DOI: 10.11646/zootaxa.3973.2.1
039A87C6C070FFF5FF3AF94BFE70FEE1.taxon	discussion	Discussion. Winterbottom et al. (2014 b) provided a BOL analysis of the CO 1 gene based on four specimens identified as T. RW sp 32 from three localities. The results suggested that there were three haplogroups corresponding to the three localities, each group separated by a minimum of 7 % of the genome from the others (Cendrawasih Bay, Group 1; Palau, Group 2; and Rabaul, Group 3). Additional localities for which no tissue samples were available include other islands within Indonesia (Bali, Sulawesi), the Philippines (several localities), the Hermit Islands and the Wallis and Futuna Islands (photograph only). A specimen from Sulawesi (Fig. 17 A) is basically similar to the Cendrawasih Bay material, but the dorsal part of the body is partly translucent, giving it a mottled appearance, the bars on the cheek are much lighter red, and the area around the bases of the first three spines of the first dorsal fin is white (which continues dorsally to join the white ocellus around the dorsal black spot). A specimen photographed alive from Bali (Fig. 17 B) has a brick-red body, with a dark grey-purple head. In two photographed specimens from Rabaul (Fig. 17 C), the background colouration of the head is a whitish-grey (rather than dark purple) heavily invested with large dark chromatophores, and there is a red blotch at the posteroventral margin of the black ocellated spot in the first dorsal fin. No basal stripe is visible in either the dorsal or the anal fins. A specimen from Palau (Fig. 17 D) has a dark grey head, with no trace of the white ocellation around the black dorsal spot, but the dark basal stripe is evident in the second dorsal fin. Specimens from the Philippines also tend to have a grey head and yellow body, but the white ocellation is clearly apparent, although one specimen has a brownish (rather than yellow) body dorsally, and the margins of the scales are red instead of black (Fig. 17 E). A specimen from the Wallis Islands (Fig. 17 6 F) is very similar to the type material, although the large dark chromatophores on the body extend further posteriorly and ventrally, and the scale margins are edged with red over most of the body. No clear-cut differences in morphometric or meristic values between the Cendrawasih Bay specimens and those from other areas were apparent. However, three of eight specimens from the Philippines examined had one or both fifth pelvic fin rays branched, as did the single specimen from Bali and one from Palau. Further work on these aspects in conjunction with wider geographic sampling of tissues is needed to explore whether the differences in CO 1 found by Winterbottom et al. (2014 b) are supported by morphology, and to investigate whether there is any genetic connectivity between the various populations / species. Specimens examined for this study were collected between 5 – 71 m. The type series collected from Cendrawasih Bay were only found between 20 – 71 m depth (and never shallower). The habitat in Cendrawasih was consistently steep outer reef walls, with T. trioculatum often present in groups of up to 20 individuals loosely aggregated and perched vertically on the substrate at the back of small overhangs and crevices. Non-type specimens collected in Bali and North Sulawesi also were also nearly always found on steep outer reef walls in the same habitat as above.	en	Winterbottom, Richard, Erdmann, Mark V., Dita Cahyani, N. K. (2015): New species of Trimma (Actinopterygii, Gobiidae) from Indonesia, with comments on head papillae nomenclature. Zootaxa 3973 (2): 201-226, DOI: 10.11646/zootaxa.3973.2.1
