A revision of the Australian species of Trimma (Actinopterygii, Gobiidae), with descriptions of six new species and redescriptions of twenty-three valid species Author Winterbottom, Richard Author Hoese, Douglass F. text Zootaxa 2015 3934 1 1 102 journal article 10.11646/zootaxa.3934.1.1 0abffe3e-a2c6-4623-b7c6-c8b26cee4a2b 1175-5326 236066 11C2A2CB-30B3-4694-B379-AE9D47332F0C Trimma lantana Winterbottom & Villa, 2004 Lantana Pygmygoby Figs 18 , 21–23 , Pl. 2 B Trimma lantana Winterbottom & Villa, 2004:16 (Honiara, Guadalcanal, Solomon Ids); Allen & Erdmann, 2012 :940 ( Indonesia to Solomons, Australia ). Trimma caesiura : Kuiter & Debelius, 2006 :662 ( Indonesia ) . Trimma sp. 1: Kuiter, 1992 :236 . Trimma sp.: Burgess et al. , 1990 :577 . Australian Material. Queensland. Cape Melville, reef off: AMS I.20774-116, 2(19–21), 0– 8 m . Escape Reef: AMS I.22575-045, 2(19–20), 7–9 m ; AMS I.22578-060, 3(20–22), 6–10 m ; AMS I.22581-044, 7(16–21), 10–14 m ; AMS I.22613-045, 2(21–23), 27 m ; AMS I.22633-082, 9(17–24), 2–11 m ; AMS I.22639-029, 2(20–21), 9 m ; ROM 40539, (22), 3–11 m ; ROM 40611, (24), 3– 15 m . Lizard Island : AMS I.18739-079, 5(17–23), 3–10 m ; AMS I.18755-109, (16), 4–8 m ; AMS I.19108-086, 15(7–24), 1–11 m ; AMS I.20730-010, 5(13–20), 20– 22 m . AMS I.20783-008, 2(21–22), 2–4 m ; AMS I.20994-007, (19), 2–7 m ; AMS I.22732-009, (19), 0–1 m ; NTM S.11444- 0 0 1, (25); ROM 1172CS, 2(19–19), 3–15 m ; ROM 39328, 2(20–21), 3– 15 m . McGillivray Reef: AMS I.19482- 133, 7(9–21), 3– 25 m . One Tree Id: AMS I.20212-021, (18), 12– 15 m . Raine Id: AMS I.20775-122, 2(22–23), 0– 20 m . Tijou Reef: AMS I.20779-143, 11(11–22), 0–25 m ; AMS I.20956-019, 15(15–27), 22–28 m ; ROM 40551, 3(22–22), 3– 12 m . Yonge Reef: AMS I.18740-074, 6(8–22), 9–12 m ; AMS I.19456-072, 4(17–20), 5–15 m ; AMS I.19481-041, (18), 2–6 m ; AMS I.20784-021, 19(15–21), 1–15 m ; ROM 39333, 2(16–19), 8– 13 m . Timor Sea off Western Australia : Ashmore Reef: NTM S.12307-022, (17), 18–20 m ; NTM S.12318-057, 2(19–22), 14–16 m ; NTM S.12322-048, (20), 12–13 m ); WAM P.29044-056, (19), 15–18 m ; WAM P.29047-034, 2(21–23); WAM P.29047-034, 2(21–26), 11– 16 m . Cartier Reef: NTM S.12883-080, 3(12–22), 13–14 m NTM S.12884-012, (22), 12– 13 m . Hibernia Reef: NTM S.13427-026, (27), 25 m . Western Australia : North Reef: NTM S.11383-029, (26), 20 m . North West Cape: WAM P.25639, (25), 8–10 m ; WAM P.26631-040, 7(18–29), 12– 14 m . Point Quobba: WAM P.27970-037, 5(17–21). Scott Reef: AMS I.21316-043, 2(18–19), 7–10 m ; NTM S.11376-040, 2(24–25). Other material. Indonesia : Ambon : USNM 209773, 3(16–21), 2–3 m ; USNM 210256 , (20), 0–8 m ; USNM 298753, 6(16–22), 14– 16m . Banda: USNM 264583, 11(15–24), 0– 3 m . Liang: USNM 209664, (17), 8 m . Kai Ids: USNM 262643, (17). Raja Ampat Ids: ROM 84901, (24), 8–25 m ; ROM 84929, 6(12–25), 2–5 m ; ROM 85115, (19), 3–7 m ; ROM 85158, (20), 20–23 m ; ROM 85231, 3(20–23), 12–16 m ; ROM 85284, 4(9–23), 2–28 m ; ROM 85323, 2(18–21), 18–21 m ; ROM 85336, 7(9–22), 3 m ; ROM 85375, 2(18–20), 14–18 m ; ROM 87435, 2(19–21), 18–24 m ; ROM 87449, 2(22–24), 15–18 m ; USNM 258775, (15), 0–6 m ; WAM P.31541-006, 5(9–23), 18–23 m ; WAM P.32255-010, 5(14–18), 8–14 m ; WAM P.33021-019, (22), 12– 18 m . Palau : Helen Reef: ROM 83077, 11(9–23), 2–8 m ; ROM 83104, 4(18–22), 7–23 m ; ROM 83136, 9(12–22), 4–8 m ; ROM 83147, (16), 8–20 m ; ROM 83193, (18), 14–23 m ; ROM 83219, 4(20–22), 15–28 m ; ROM 83233, 5(15–19), 8– 20 m . Papua New Guinea : Hermit Ids: USNM 263623, 2(8–13), 0–8 m ; USNM 264517, 10(14–25), 0–12 m ; USNM 264544, 4(17–19), 0–15 m ; USNM 264556, 13(12–24), 0–12 m ; USNM 264559, 6(18–23), 0– 33m . Louisiade Islands: WAM P.31736-020, 2(21–22), 20– 25 m . Manus Id: WAM P.27827-046, (13), 7– 9 m . Ninigo Ids: USNM 298775, (12). Port Moresby: AMS , (21), 18 m . New Britain : ROM 92115, 2(15–16), 3 m . Trobriand Ids: USNM 293966, (21), 0–8 m ; USNM 295524, (21). Solomon Islands : Hope Id: NTM S.12710-008, (19), 6 m . Florida Island : AMS I.17499-022, (17), 8– 12 m . Guadacanal: BPBM 16159, (19), 10 m ; ROM 46039, (20.8), 21 m ; ROM 46044, 2(17–19), 5 m ; ROM 73436, 5(12–21), 21 m . Savo Id: AMS I.17491-038, (17), 10 m . Diagnosis. A species of Trimma with a deep interorbital trench, a deep groove behind the upper half of the eye; second spine of first dorsal fin longest, elongated into a short filament, third sometimes filamentous; dorsal and anal rays usually I 8 ; midline of predorsal region completely covered with ctenoid scales in 5–8 rows, reaching anteriorly to the interorbital trench; operculum usually naked, but sometimes with one or two small scales dorsally; pectoral-fin base with two vertical rows of 4–5 cycloid scales each, uppermost 1–2 scales largest (distinctly enlarged); prepelvic area covered with 5–8 rows of cycloid scales; upper and lower pectoral rays unbranched, central 5–10 rays branched; pelvic fins largely separate, connected only at the base with one or two scales on the midline extending beyond the connecting membrane, left and right fins widely separated, distance between the bases of the fins about half to three–quarters of the base of each fin; fifth pelvic ray usually with a single dichotomous branch, others with a single sequential branch (2 terminal tips); fifth ray 50–90% the length of the fourth; nape crest absent; cheek with a bar below the anterior part of the eye; head with large reddish to brown spots (Pacific) or vertical bars (Western Australia ), slightly narrower than the pupil diameter; pectoral-fin base usually with 3 reddish spots (light areas in preserved material); body with two zigzag lines anteriorly on the body, large white spots posteriorly on the caudal peduncle and below the bases of the second dorsal and anal fins; and caudal peduncle lighter than the anterior part of the body (yellow in life). Description. The description is primarily based on specimens from the Great Barrier Reef and Solomon Islands . Dorsal fin usually VI + I 8 ( I 7 or I 9 in 4.8% of individuals), third spines longest, elongated into filament (reaching first segmented rays in juveniles, reaching well beyond end of second dorsal fin in some large adults); first ray of second dorsal fin usually unbranched (branched in 15–30% of individuals in any sample); anal fin I 8 (rarely I 7 or I 9 , in 6.1% of individuals), first ray usually unbranched (branched in 15–30% of individuals in any sample); pectoral-fin rays 15–18 ( 16 in 66% of specimens, 17 in 26% of specimens), reaching posteriorly to vertical in line from just beyond anal spine to about second segmented ray; pelvic fin I 5 , fins widely separate with very low basal membrane, which is covered by a median scale, distance between bases of fins slightly less than base of each pelvic fin, first four rays with one sequential branch, fifth ray with one dichotomous branch (two terminal tips) or unbranched, 50–90% length of fourth ray, which reaches posteriorly to below point between anus and anal spine; segmented caudal rays usually 17; branched caudal rays usually 6 upper + 5 lower (5 + 5 or 6 + 6 in <3% of specimens). Lateral scales 22–24, usually 23; transverse scales 6–8, usually 7; predorsal scales on midline 5–8, usually 6–7, anterior-most of which sometimes distinctly larger than scale behind it, but often subequal in size, scales extending to edge of postorbital trench above posterior part eye; preoperculum scaleless, operculum scaleless or with 1–2 small cycloid scales; scales below second dorsal fin with small accessory scales; pectoral-fin base with two vertical rows of 4–5 large cycloid scales, scales ovoid; prepelvic area with 6–8 rows of cycloid scales; remaining scales ctenoid. Teeth in lower jaw consist of an outer row of curved, slightly enlarged canines ending near bend in dentary and inner row of similar, but smaller canines extending full length of dentary, with irregular row of small conical teeth in between; outer row of teeth in upper jaw similar to those of lower jaw, with 1–2 small irregular inner rows of teeth, innermost row not enlarged. Tongue tip variable in shape, usually with pointed triangular anterior margin, rarely rounded or bilobed. Gill opening extending anteroventrally to below posterior margin of pupil; outer gill rakers on first arch very elongate, almost equal to filament length at angle 2–5 + 15–18 = 19–23, n = 50, mean = 3.6 + 16.4 ( Winterbottom & Villa, 2003, reported 3–4 + 15–17, n = 6, mean = 3.4 + 15.7 ). Anterior nares a narrow tube, posterior nares small pore with raised rim; nasal sac elevated as raised oval sac located just above upper lip, openings separated by about 4–5 nares diameters. Interorbital as described in diagnosis (see also Fig. 1 A of T. caesiura ), epaxialis musculature attaching to posterior wall of interorbital/postorbital trench. Abdominal/caudal vertebral transition is Type B. FIGURE 21. Trimma lantana . A: AMS I.22633, Escape Reef, Queensland; B: North-West Cape, Western Australia; C: Halmahera, Indonesia. Photos by D. F. Hoese (A) and G. R. Allen (B & C). Colour pattern. Freshly collected specimens from Pacific. (From slides of freshly collected specimen from Yonge Reef and Escape Reef ( Fig. 21 A), Queensland, Raja Ampat Ids, Papua , Guadalcanal, Solomon Ids and Helen Reef, Palau ). Head with large red spots, bordered by thin black rim; anterior spots often dark brown obscuring red colouration; interspaces white forming and irregular network of interconnected lines; snout usually dark blue to dark grey; thin vertical bar below anterior margin of pupil extending to near posterior end of jaws; vertically elongate dark spot below middle of pupil, not reaching to eye, varying in width from thin line to almost as wide as pupil diameter; large oval spot just below posteroventral margin of eye, sometimes triangular or slightly vertically elongate; below this spot vertically elongate to oval reddish spot, slightly smaller than spot in contact with eye; two reddish vertically elongate spots along posterior preopercular margin, often crossing onto operculum, rarely connected to form hourglass-shaped mark; operculum usually with two large spots, one dorsal and one ventral; small red spot directly behind posterodorsal margin of eye, followed dorsally by large red spot; nape with very large (almost equal to eye diameter) triangular spot; white interspaces from head extending onto body as two zigzag lines, one below dorsal fins and one just below middle side; lower stripe sometimes with vertical extension to anal fin base and sometimes other vertical extensions reaching to near anal fin; posteriorly both white lines on body breaking into small white spots; body reddish anteriorly, yellow to orange on caudal peduncle, yellow varying from covering whole of caudal peduncle to only posterior area just before caudal fin; pectoral-fin base with 3 oval red spots; one anteriorly on middle of pectoral-fin base and two posteriorly, one above and one below; upper spot with small mark posterodorsally, sometimes extending ventrally as thin line; dorsal fins with small spots proximally forming horizontal stripe near base of fin; second dorsal fin with small reddish to yellow spots forming 2–3 horizontal lines above base of fin, fin becoming paler distally; anal fin largely clear to dusky, often with small red spots near base of fin; caudal fin with yellow to orange spots often forming wavy bands, more prominent basally; pelvic fin dusky to white, sometimes with faint yellow or orange indistinct markings. Colouration in life . (Based on a photo by A. Gonzalez-Cabello of an individual from Lizard Island , Queensland, Pl. 2 B). Similar to colouration of freshly collected specimens, except that red colouration more intense and caudal peduncle translucent yellow. Winterbottom & Villa (2004) give detailed description of live and freshly collected material from the Great Barrier Reef. Freshly collected specimens from mainland of Western Australia . ( Fig. 21 B). Head with vertical bands and vertically elongate red to orange spots, bordered by thin black rim; a broad (slightly narrower than pupil diameter) vertical bar below anterior margin of pupil extending to near posterior end of jaws; second vertical bar from posteroventral margin of eye to lower preoperculum; snout slightly darker than rest of head; two reddish orange vertically elongate spots along posterior preopercular margin, often crossing onto operculum; operculum with one large spot dorsally and sometimes faint grey mark ventrally; small orange spot directly behind posterodorsal margin of eye, followed dorsally by large red spot; nape with very large (almost equal to eye diameter) triangular spot; large white interspaces from head extending onto body as two zigzag lines, one below dorsal fins and one just below midside; lower stripe with vertical extension to anal fin base and other vertical extensions reaching to near anal fin and ventrally on caudal peduncle; posteriorly both white lines on body breaking into large white spots; body reddish orange anteriorly, yellowish-orange on caudal peduncle, but only slightly lighter than anterior colouration; pectoral-fin base with three oval red spots; one anteriorly on middle of pectoral-fin base and two posteriorly, one above and one below; upper spot large with wide dark bar posteriorly; fin colouration as described above. Freshly collected specimen from Halmahera . ( Fig. 21 C). Head with vertical bands and vertically elongate red spots, bordered by thin black rim; broad (about half pupil diameter) vertical bar below anterior margin of pupil extending to near posterior end of jaws; short second vertical bar from posteroventral margin of eye to middle of preoperculum; snout slightly darker than rest of head; two reddish orange vertically elongate spots along posterior preopercular margin, crossing only slightly onto operculum; operculum with one large red spot dorsally and faint grey mark ventrally; small orange spot directly behind posterodorsal margin of eye, followed dorsally by slightly larger red spot; nape with large (equal to pupil diameter) round spot; large white interspaces from head extending onto body as two zigzag lines, one below dorsal fins and one just below midside; lower stripe with vertical extension to anal fin base and other vertical extensions reaching to near anal fin and ventrally on caudal peduncle; posteriorly both white lines on body breaking into large white spots; body red orange anteriorly, yellow on caudal peduncle, but distinctly lighter than anterior colouration; pectoral-fin base with three oval red spots; one anteriorly on middle of pectoral-fin base and two posteriorly, one above and one below; upper spot large with two small black spots posteriorly; fin colouration as described above. Preserved . Snout dark brown; head with large dark brown spots or bands below eye, corresponding to red spots in fresh material; spots on preoperculum margin and operculum dark edged with light brown in center; overall body colour light brown; scales edged in melanophores more prominently anteriorly; entire head and body very faded in material from Timor Sea; fins clear, with scattered melanophores, but no distinct spots; pectoral-fin base light brown, sometimes with traces of three spots, but distinct dark bar on pectoral-fin base in specimens from mainland of Western Australia . Variation. The species is most similar to Trimma caesiura and T. naudei . Winterbottom & Villa (2004) compared it with T. naudei and other relatives. They only reported colour differences between Trimma lantana and T. naudei , separating the former from the latter by the absence of a dark, vertical pectoral bar and the presence of two separate orange spots on the vertical limb of the preopercle (versus a club-shaped bar). As noted below, the absence of the dark bar on the pectoral-fin base does not hold up for mainland Western Australian specimens. In addition, the bar or spots on the vertical limb of the preopercle are variable in Timor Sea material. We note that, for all forms of T. lantana , the caudal peduncle is lighter than the body in freshly collected material (versus caudal peduncle the same colour as the body in T. naudei ). We also note here a statistical difference in the presence of an opercular scale. In over 100 well-preserved specimens from the Great Barrier Reef only 4 specimens were found with the scale. In 25 of 40 T. naudei examined the scale is present. Furthermore, specimens of T. lantana from Western Australia more commonly have the scale as well. Many of the specimens from Western Australia are not in good condition and we cannot be sure of the exact occurrence (but did find the scale in about one-third of the specimens). FIGURE 22. Head pigmentation patterns in Trimma naudei . A: Philippines, USNM 263469; B: Madagascar, WAM P.32022- 0 12. Trimma lantana differs consistently from T. naudei in having spots or bands below the eye and a paler caudal peduncle. However, it appears that there may be several species in the T. naudei complex based on colour patterns. Trimma naudei , as currently recognised, may actually consist of two species, one in the Indian Ocean and one in the Pacific. Specimens from the Pacific have a reddish body with large white areas, with the caudal peduncle the same colour as the body, and there is a dark area on the pectoral-fin base in the form of a one-third pupil diameter wide vertical line ( Fig. 22 A).The typical T. naudei from the Indian Ocean the body colouration is the same as specimens from the Pacific Ocean, but there is a dark oval spot almost pupil diameter in width dorsally on the pectoral-fin base rather than a vertical line ( Fig. 22 B). As recognised here, three different colour phases of T. lantana are known from Australia . The form found on the Great Barrier Reef ( Fig. 23 A), Solomon Islands , Papua New Guinea and Papua , Indonesia ( Fig. 23 B) has distinct dark spots under the eye and three light areas on the pectoral-fin base. Specimens from the mainland of Western Australia ( Fig. 23 C and Fig 21 B), are similar in body colouration, with a light caudal peduncle, but differ in having dark vertical bars under the eye and a prominent dusky bar at the base of the pectoral fin. In some specimens the posterior band on the cheek is only a vertically elongate spot, not reaching to the lower edge of the preoperculum. The bar at the base of the pectoral fin is darker and broader than the spot or bar in either form of T. naudei . Specimens from Scott Reef ( Fig. 23 D), which lies well offshore, are lighter and only have faint traces of bands under the eye and a faint bar on the pectoral-fin base. The third form is known from Timor Sea islands and parts of Indonesia (Ceram and Halmahera). The live colouration of material from the Timor Sea is unknown, but preserved material usually has one spot posteriorly on the cheek below the eye ( Fig. 23 E–F), similar to freshly collected specimens from Halmahera ( Fig. 21 C). In some specimens from the Timor Sea ( Fig. 23 E) there are traces of faint spots below the middle of the eye, but these are never as prominent as in specimens from the Pacific. Some specimens show faint trances of bands under the eye and some have a faint vertical bar on the pectoral-fin base. Specimens from Ashmore and Cartier Reefs are highly variable and it is possible that more than one species is present on those reefs. Without live colouration, it is not possible at present to separate the various forms found on these reefs. Alternatively the area may represent a transition area between T. naudei and T. lantana with some interbreeding. Consequently without genetic information and life colouration of the Ashmore Reef and surrounding areas we recognise all of the Australian material as T. lantana , but realise that two or more species may be involved. FIGURE 23. Head pigmentation patterns in Trimma lantana . A: Lizard Id, Great Barrier Reef, AMS I.19108–086; B: Raja Ampat Ids, Papua, WAM P.33021–019; C: mainland, Western Australia, WAM P.27970-037; D: Scott Reef, Western Australia, NTM S.11376–040; E: Ashmore Reef, Timor Sea, NTM S.12318-057; F: Cartier Reef, Timor Sea, NTM S.12883-080. Photos by D. F. Hoese. Analysis of variance of the lower gill-raker counts of the various populations (Great Barrier Reef and Solomon Island , n = 50; Western Australia to Halmahera, n = 23; Trimma naudei from the western Indian Ocean (n = 27) and T. naudei from Philippines and Japan , n = 20), indicates a significant difference in counts (p <.05), with counts averaging 16.4 for T. lantana ( Australia and Solomon Islands ) and 16.3 (Western Australia to Halmahera); 14.5 for T. naudei (western Indian Ocean) and 15.5 ( Philippines and Japan ). Although the sample sizes are too small for most populations to make individual comparisons, the differences between the two forms of T. naudei are as great as the differences between Pacific T. naudei and T. lantana . Table 2 gives the raw data for lower gill raker counts. TABLE 2. Lower gill raker counts for Trimma lantana and T. naudei .

Lower gill raker count	13	14	15	16	17	18
Australia - Solomon Islands	-	-	4	27	16	3
Western Australia to Halmahera	-	1	5	6	9	2
Western Indian Ocean	3	10	13	2	-	-
Philippines and Japan	-	3	7	8	2	-

Etymology. Named after the tropical plant Lantana , in allusion to its red and yellow flowers that are reminiscent of the predominant colours of this species. The name is a noun in apposition. Distribution. Trimma lantana is known from mid and outer-edge reefs of the Great Barrier Reef and from Western Australia . The species is not known from inshore reefs on the Great Barrier Reef. Provisionally we include material here from offshore reefs in the Timor Sea and islands and mainland areas off Western Australia as Trimma lantana (see following discussion). The species is known from depths of 1– 28 m . Elsewhere, the species has been recorded from eastern Indonesia , Papua New Guinea , the Solomon Islands and from Helen Reef, Palau (but is apparently absent from the main Palauan islands). Comparisons. For the differences between T. lantana and other members of the T. caesiura group, see under T. caesiura (above). Results from an analysis of the CO1 gene ( Winterbottom et al. 2014 ) included seven specimens of T. lantana (Great Barrier Reef, n = 2; Helen Reef, Palau , n = 2, Raja Ampat, Indonesia , n = 2; and Rabaul, New Britain , n = 1) in a single haplogroup, with a within group variation of 0.8%. Specimens of T. naudei from the western Indian Ocean (n = 7) were separated from specimens from Japan and Taiwan (n = 3) by a minimum of 3%, and these two groups differed from the T. lantana specimens by between 2.3% (western Indian Ocean) and 4.0% ( Japan / Taiwan ). No other specimens identified as T. naudei were available for that study, and it would be especially interesting to obtain specimens from Western Australia , central Indonesia , the Philippines and Vietnam for analysis. Discussion. This species has been referred to informally as Trimma DFH sp. 13 or T . RW sp. 56. It reaches a maximum recorded size of 29.4 mm SL.