A new species of Trimma of the T. taylori species group (Teleostei: Gobiidae) from the Red Sea, Indian Ocean.

A new species of Trimma is described from the Red Sea along the Saudi Arabian coast. Specimens and/or photographs of this species are available from the Egyptian Red Sea to Eritrea. These specimens, formerly identified as T. taylori, differ from all other samples from the Indo-Pacific currently identified as T. taylori in having 9 and 89 dorsal- and anal-fin rays respectively (vs. usually 10 and 10 rays), 13 pectoral-fin rays (vs. usually 14 rays), and cycloid scales covering the entire predorsal region from the upper base of the pectoral fin anterior to a convex line posterodorsally to just lateral to the base of the sixth first dorsal-fin spine (vs. predorsal region mostly or entirely covered with ctenoid scales). In addition, specimens from the Red Sea form a monophyletic lineage in a Maximum Likelihood analysis of the COI gene. In this tree, the new species is the sister group to a clade composed of three lineages. One is composed of specimens from the Maldives, which is the sister group of a single available specimen from the Seychelles. These two together are the sister group of specimens of a widespread western Pacific clade.

A new species of Trimma (Pisces: Gobiidae) from the deep reefs of Palau, western Pacific Ocean.

A new species of Trimma is described from three specimens from deep reefs (91.4 m) at Uchelbeluu Reef, Palau, western Pacific Ocean. Trimma panemorfum n. sp. is characterized by a live colouration of a yellow to orange body with two light blue stripes, each with a ventral bar of the same colour from the anterior origin. The predorsal midline is scaled, opercular and cheek scales are absent, the middle 1213 pectoral-fin rays are branched, the fifth pelvic-fin ray has two dichotomous branch points (total of four branch tips), the bony interorbital is 3442% pupil width and does not extend ventrolaterally beyond the fifth papilla of row p, where the posterior interorbital trench is present as a slight groove or absent.

A new species of Trimma (Pisces: Gobiidae) from western Thailand, north-eastern Indian Ocean.

A new species of Trimma is described from the islands just off the Thailand coast near Phuket. Trimma ukkriti n.sp. lacks scales on the cheek and opercle, but has ctenoid scales in the midline of the nape, has an elongate second spine of the first dorsal fin which reaches posteriorly to between the base of the spine and the base of the 4th ray of the second dorsal fin when adpressed, 19-20 pectoral-fin rays with 6-13 branched rays in the middle of the fin, an unbranched 5th pelvic-fin ray, which is 48-67% the length of the 4th ray, a U-shaped interorbital trench with no fleshy median ridge, a poorly developed postorbital trench ending at the 5th papilla in row p (which has 6 papillae in total), and row c beneath the eye consists of 6 papillae. When live and freshly collected, the species is very similar in appearance to T. okinawae and T. readerae, with orange to red spots on the head and body.

A new species of Trimma (Pisces: Gobiidae) from the northern Palauan Islands, western Pacific Ocean.

A new species of Trimma is described from the northern Palaun islands. Trimma kalum n. sp. lacks scales on the cheeks, opercle and nape, has an elongate second spine of the first dorsal fin, reaching posteriorly to the base of the 2nd-10th ray of the second dorsal fin when adpressed, 17-18 pectoral-fin rays with 12-13 branched rays in the middle of the fin, a single dichotomous branch point in the 5th pelvic-fin ray, which is 51-75% the length of the 4th ray, a very well developed dermal crest in the midline between the base of the first dorsal spine and the posterior interorbital region, a U-shaped interorbital trench with no fleshy median ridge, a poorly developed postorbital trench ending between the 4th to 5th papillae in row p, which has 6 papillae in total, and there are 5 papillae in row c beneath the eye. When live and freshly collected, the species is very similar in appearance to T. sheppardi, with an overall yellow body, three yellow bars across the cheek, and a double dark spot above the posterior portion of the opercle.

A new species of Trimma (Pisces; Gobiidae) from the Western Pacific Ocean.

A new species of Trimma, T. wangunui, is described from three localities in the western Pacific (Papua New Guinea, Timor-Leste and the Philippines). It belongs to a small group of species with scales in the predorsal midline, no scales on the cheek or the opercle, all pectoral fin rays unbranched, and a branched fifth pelvic fin ray. It differs from other species in this group in having an elongate second spine of the first dorsal fin which reaches to the bases of the 2nd-8th second dorsal-fin rays when adpressed, in having yellow bars on the head, and in the presence of vertically elongate yellow spots on a brown body when freshly collected.

Two new species of blue-eyed Trimma (Pisces; Gobiidae) from New Guinea.

Two new species of Trimma are described from New Guinea, one at the southeastern end at Normanby Island (Milne Bay Province), the other from Cendrawasih Bay, West Papua, on the north-east coast. The dorsal surface of the eye of both species is blue in life, a characteristic not reported elsewhere in the genus. Although the two species look very similar in life, and both occupy similar mesophotic rubble habitats in the 50-70 m depth range, they are separated both genetically (7.7% pairwise genetic distance in COI) and morphologically. Trimma blematium has 16 pectoral fin rays, a branched 5th pelvic fin ray, and 7 papillae in row p, whereas T. meityae has 17-18 pectoral fin rays, an unbranched 5th pelvic fin ray, and 8 papillae in row p. In live specimens, the blue colour over the top of the eyes is much darker in T. blematium than in T. meityae. The type localities are separated by almost 2,000 km (straight-line distance).

A new species of Trimma (Pisces; Gobiidae) from the South-West Islands of Palau, western Pacific Ocean.

A new species of Trimma is described from South-West Islands of Palau. Trimma hamartium n. sp. lacks scales on the cheeks and opercle, has 8-9 scales in the predorsal midline, 17-19 pectoral-fin rays with 5-10 branched rays in the middle of the fin, an unbranched 5th pelvic-fin ray that is 51-64% the length of the 4th ray, 17-19 gill rakers on the outer surface of the first gill arch, an U-shaped interorbital trench with a narrow, slit-like postorbital trench ending at the last papilla in row p, and has 6 papillae in row c beneath the eye. When freshly collected, the species is very similar in appearance to T. preclarum, with which it is syntopic at three of the more northern islands of the South-West Islands.

Two new species of Trimma (Pisces; Gobiidae) from Fiji, south-western Pacific Ocean.

Two new species of Trimma are described from Fiji. Trimma bathum n. sp. lacks scales on the cheeks, opercle and predorsal midline, has 18-19 unbranched pectoral fin rays, an unbranched 5th pelvic fin ray that is 40-56% the length of the 4th ray, 17-18 gill rakers on the outer surface of the first gill arch, a U-shaped interorbital and a narrow slit-like postorbital trench, a low, median fleshy ridge extending half-way towards the orbit from the origin of the first dorsal fin, and, when freshly collected, a pink head and body with most body scales having an orange-brown spot or short bar at their centres. The species is currently known only from off Suva Harbour, Viti Levu, Fiji. Trimma finistrinum n. sp. has a bony interorbital equal to the pupil diameter, a fully scaled nape of 12-14 scales, a second dorsal spine that may reach posteriorly to the middle of the second dorsal fin, the papillae in the longitudinal row immediately below the eye either single or with two papillae in a vertical row, unbranched pectoral fin rays, usually a branched fifth pelvic-fin ray that is about half length of the fourth ray, and a large diffuse dark blotch on the posterior part of the caudal peduncle. A colour pattern of a brownish body with most body scales having golden- to greenish-yellow (pale in preservative) centres is unique among species of the genus. The species is currently recorded only from off the north and east coasts of Viti Levu, Fiji.

Trimma tevegae and T. caudomaculatum revisited and redescribed (Acanthopterygii, Gobiidae), with descriptions of three new similar species from the western Pacific.

A redescription and diagnosis of Trimma tevegae, based on 50 additional specimens from the type locality at Rabaul, New Britain is provided, and contrasted with the redescription and diagnosis of T. caudomaculatum, based on the type material and an additional 22 specimens from various western Pacific localities. Trimma tevegae may most easily be recognized by the lack of a blue stripe (dark in preservative) in the dorsal midline of the snout, and a short second spine in the first dorsal fin, reaching posteriorly to the origin or first few ray bases of the second dorsal fin. Trimma caudomaculatum can be identified by the blue stripe from the snout along the dorsal midline, the blue spots and stripes in front of the eye, on the opercle and beneath the eye (dark in preserved material), and the very elongate second dorsal spine, usually extending to beyond the end of the second dorsal fin. Three similar-looking new species are described. Trimma burridgeae and T. hollemani are morphologically very close to each other, differing mainly in the length of the second dorsal spine (to the posterior base of the second dorsal fin or beyond in T. burridgeae; to the anterior base of that fin in T. hollemani). These two species differ by over 9% of the bases of the COI gene. Trimma corerefum is the most distinctive of the species, differing in lacking a blue stripe on the dorsal surface of the snout, in the pigment pattern around the eye, in having fewer cephalic sensory papillae (free neuromasts) in rows d', ea, ep, f, r and ot, and in a shorter fifth pelvic fin ray (34-45% length of fourth pelvic ray). The morphological differences between the species are reinforced by the results of a barcode analysis of the COI gene, based on 105 specimens.

New species of Trimma (Actinopterygii, Gobiidae) from Indonesia, with comments on head papillae nomenclature.

Three new species of the gobiid Trimma are described from Indonesian waters, and a partially reformulated nomenclature for the cephalic sensory papillae of members of this genus is provided. Trimma aturirii possesses 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, with the dorsal half of the body 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, 17-18 (7-11 branched) pectoral fin rays and an unbranched fifth pelvic fin ray. Trimma kardium has a pair of tapering oval red spots which join anteriorly over the anterior region of the hyoid arches, forming an approximate heart-shaped marking on the ventral surface of the head. It has 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 nape, and a single row of 1-3 cycloid scales along the upper border of the opercle. Trimma trioculatum has 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.

A revision of the Australian species of Trimma (Actinopterygii, Gobiidae), with descriptions of six new species and redescriptions of twenty-three valid species.

The gobiid genus Trimma currently contains 75 valid species, with another 20-30 known but undescribed species. There are 29 species in Australian waters (six undescribed). This paper describes the six new species, and provides redescriptions of most of the 23 previously described species known from the region, as well as a key for all the species. The six new species are: T. insularum (endemic to Cocos (Keeling) Islands), T. kitrinum (Fiji to Great Barrier Reef), T. meristum (Cape York to the Bismark Archipelago and Fiji), T. pentherum (Great Barrier Reef to Fiji and the South-West Islands of Palau), T. readerae (Australia to Japan), and T. xanthum (Palau to Fiji, Great Barrier Reef to Christmas Island). The following 23 species have been recorded from Australian waters, and most are redescribed here: T. anaima (Comores to Fiji), T. annosum (Maldives to the Phoenix Islands, Taiwan to the southern Great Barrier Reef), T. benjamini (southern Vietnam to the Marshall Islands, Samoa and southern Barrier Reef), T. caesiura (Ryukyus through the Marshall Islands to Samoa and Elizabeth Reef on the Lord Howe Rise), T. capostriatum (New Caledonia to eastern Australia and Papua New Guinea), T. maiandros (Java to the Ryukyus, Marshalls to Great Barrier Reef), T. emeryi (Comores to Ryukyus and Samoa), T. fangi (western South China Sea through to the Solomons), T. flavatrum (Ryukyu Islands to Western Australia and Samoa), T. hoesei (Chagos Archipelago, central Indian Ocean to Palau and Solomons), T. lantana (Australia, Solomons, northern New Guinea, South-West Islands of Palau), T. macrophthalmus (Ryukyu Islands to Cocos (Keeling) Islands and Samoa), T. milta (Taiwan to Western Australia, Society Islands and Hawaii), T. nasa (Sumbawa, Indonesia to Fiji), T. necopinum (northern tip of Cape York to Sydney), T. nomurai (Japan to northern Australia and New Caledonia), T. okinawae (western Thailand to Japan and the Phoenix Islands, north-west Australia to the Great Barrier Reef), T. preclarum (Palau to Fiji, Great Barrier Reef), T. stobbsi (Maldives to New Caledonia), T. striatum (Maldives to Palau, to northern Australia), T. taylori (Red Sea to Hawaii and Society Islands), T. tevegae (Red Sea to Ryukyu Islands, Marshall Islands to Samoa), and T. unisquame (Comores to Hawaii and Easter Island).

Three new species of Trimma (Pisces; Gobioidei) from Indonesia.

Three new species of Trimma are described from various localities in Indonesia. All three can be readily identified from their live, freshly collected, or preserved colouration. Trimma meranyx n. sp. is further distinguished from other species by the possession of 8-9 scales in the predorsal midline, up to three rows of (usually) cycloid scales on the opercle, two scales at the posterodorsal border of the cheek, a very slightly elongate second dorsal spine which only just reaches the spine or anterior rays of the second dorsal fin, unbranched pectoral fin rays, a fifth pelvic fin ray that branches once and is 64-85% the length of the fourth ray, and a full basal membrane connecting the inner branches of the two fifth pelvic rays. The dark red (live) or black posterior half of the caudal peduncle with large white spots straddling the dorsal and ventral midlines just anterior to the first procurrent caudal fin rays is the diagnostic colour character. The species is known from North Sulawesi, West Papua (Raja Ampat and Fakfak), and the south-eastern tip of Papua New Guinea, with possible records from the Philippines and Vanuatu. Trimma pajama n. sp. has 6 scales in the predorsal midline, two ctenoid scales along the dorsal margin of the opercle, a slightly elongate second dorsal spine reaching posteriorly to the base of the spine or first ray of the second dorsal fin, unbranched pectoral fin rays, a fifth pelvic ray with a single branch point and which is 58-72% the length of the fourth ray, and a full basal membrane connecting the inner branches of the two fifth pelvic rays. The live, freshly collected and preserved colour pattern of alternating dark and light stripes on the head and most of the body (except the posterior half of the caudal peduncle) is diagnostic. It is currently known from West Papua (Raja Ampat and Fakfak) and the southern tip of Papua New Guinea, with possible records from Kalimantan (Indonesia), Palau, the Hermit Is (Papua New Guinea) and the Solomon Islands. Trimma zurae n. sp. has 8-9 scales in the predorsal midline, usually a single row of cycloid scales along the upper border of the opercle, 11 anterior and 9 posterior transverse scale rows, no elongated spines in the first dorsal fin, 9 dorsal and 8 anal fin rays, the middle rays of the pectoral fin branched, a single branch in the fifth pelvic fin ray which is 65-76% the length of the fourth ray and a reduced basal membrane of < 20% the length of the fifth ray. The eye-diameter sized black ocellated spot between the first to fifth spines of the first dorsal fin is diagnostic, as are the pupil-diameter sized orange spots on the nape, opercle and posterodorsal part of the cheek. It is currently known only from a single locality just west of Manado, Sulawesi. 

Eviota piperata, a new gobiid species from Palau (Teleostei: Gobiidae).

A new species of dwarfgoby, Eviota piperata is described from Palau. It belongs to the cephalic sensory-pore system Group II (lacking only the IT pore); has a dorsal/anal-fin formula of 8/8; has some pectoral-fin rays branched; no dark spot over the ural centrum; the male genital papilla is not fimbriate; and the cheek and body are heavily peppered with chromatophores.

Trimma helenae (Pisces; Gobioidei), a new species of gobiid fish from Indonesia.

A new species of Trimma, T. helenae, is described from the southeastern lagoon at Penemu Island off the southwest coast of Waigeo, Raja Ampat, Indonesia. The new species has a unique colour pattern when alive, consisting of a yellow anterior half and red posterior half, with four small white spots along the midline of the dorsal and ventral surfaces of the caudal peduncle. It is also the only species of the genus to have a nasal sac that is flush with the snout surface (not raised above the level of the snout or only represented by a nasal pit), and which lacks a raised rim to the posterior nasal pore. Trimma helenae belongs to a group of 12 valid nominal species defined by having a broad bony interorbital region (width 80-100% of pupil diameter), but differs from all of but three of these in having only cycloid scales in the midline and on the sides of the nape. The other members of the group have mostly ctenoid scales in this region.

Trimma irinae, (Pisces; Gobioidei) a new species of gobiid fish from Milne Bay Province, Papua New Guinea.

A new species of Trimma, T. irinae, is described from 65 m off Lawadi, Milne Bay Province, Papua New Guinea. This colourful new species has red-orange spots on the head and anterior trunk, with a yellow to orange body posteriorly. The first dorsal fin has two dark basal spots, and the elongated section of the second dorsal spine is bright white to pale blue in life. There are 8-9 scales in the predorsal midline, usually cycloid scales present along the upper border of the opercle in up to three horizontal rows, no cheek scales, and the middle 9-11 rays of the pectoral fin are branched. The fifth pelvic fin ray branches once dichotomously and is 52-64% the length of the fourth ray, and the basal membrane connecting the inner margins of the fifth pelvic fins rays is less than 15% the length of the fifth ray.

A cornucopia of cryptic species - a DNA barcode analysis of the gobiid fish genus Trimma (Percomorpha, Gobiiformes).

A genetic analysis of partial mitochondrial 5' cytochrome c oxidase I gene (DNA barcode) sequences of 473 specimens assigned to 52 morphological species (including four known, but not formally named, species) of the gobiid genus Trimma revealed the presence of 94 genetic lineages. Each lineage was separated by > 2% sequence divergence. Thus there were an additional 42 haplogroups recognizable as provisional candidate species given that a value of > 2% difference is typical of different species of fishes. Such a high degree of apparently different cryptic species is, in our experience, virtually unprecedented among vertebrates. These results have precipitated further morphological research in a few cases, which has uncovered subtle differences independently corroborating the genetic results. However, such efforts are limited by the dearth of traditional systematists available to undertake the necessary time-consuming, and highly detailed, morphological research. In some cases, the genetic results we present are consistent with, and confirm, minor taxonomic distinctions based on morphology and/or colour pattern. In other instances, what had been recognized as a single species consists of several genetic lineages - up to eight in, for example, what we have identified based on morphology as Trimma okinawae. The increase from 52 to 94 potential species in our sampling raises the predicted total number of species in this genus from about 110 to nearly 200 (versus the 73 valid described species currently recognized).

CLADOGRAM VERSATILITY: EVOLUTION AND BIOGEOGRAPHY OF ACANTHUROID FISHES.

A highly corroborated cladogram of acanthuroid fishes is used to explore several aspects of the biogeographic and microevolutionary events during the evolution of the group. Five events in acanthuroid evolution are documented here, which demonstrate not only the power of cladograms to provide adaptational hypotheses, but also how they can be used to frame significant questions for further research. Biogeographic analysis indicates that basal cladogenesis of the acanthuroids must have occurred prior to the Eocene (50-55 mya), because at least the basal lineages of the Acanthuridae, one of the last acanthuroid families to have evolved, were present at that time. In addition, optimization of current distributions suggests that the ancestors of the Acanthurinae, of the Acanthuridae, and of the Acanthuroidei each had an Indo-West Pacific distribution. Subsequent dispersion and/or speciation in one or more of these lineages may have been related either to the closure of the Tethys (ca 20 mya), or to colonization from the Pacific of the Caribbean/Atlantic prior to the formation of the Panamanian isthmus. Hypotheses about adaptation in acanthuroids begin with a discussion of several cases of dietary plasticity. The importance of information from fossils is illustrated in an investigation of predorsal bone evolution. Morphologically divergent and convergent tendencies in the modification of the pelvic fins concludes the examples. The morphologically most derived genus, Ctenochaetus, seems the least active taxon evolutionarily in the characters investigated.