Research Article |
Corresponding author: Richard L. Pyle ( deepreef@bishopmuseum.org ) Academic editor: David Morgan
© 2018 Richard L. Pyle, Brian D. Greene, Joshua M. Copus, John E. Randall.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Pyle RL, Greene BD, Copus JM, Randall JE (2018) Tosanoides annepatrice, a new basslet from deep coral reefs in Micronesia (Perciformes, Percoidei, Serranidae). ZooKeys 786: 139-153. https://doi.org/10.3897/zookeys.786.28421
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The new species Tosanoides annepatrice sp. n. is described from four specimens collected at depths of 115–148 m near Palau and Pohnpei in Micronesia. It differs from the other three species of this genus in life color and in certain morphological characters, such as body depth, snout length, anterior three dorsal-fin spine lengths, caudal-fin length, and other characters. There are also genetic differences from the other four species of Tosanoides (d ≈ 0.04–0.12 in mtDNA cytochrome oxidase I). This species is presently known only from Palau and Pohnpei within Micronesia, but it likely occurs elsewhere throughout the tropical western Pacific.
closed-circuit rebreather, coral-reef twilight zone, mesophotic coral ecosystems, Micronesia
The authors have explored mesophotic coral ecosystems (MCEs; coral-reef habitat at depths of 30–150 m; also referred to as the “coral-reef twilight zone”) throughout Micronesia for decades, and have utilized advanced mixed-gas closed-circuit rebreather diving operations since 1997, which has led to the discovery of dozens of new species of fishes (
The specimens represent an undescribed species of the serranid subfamily Anthiadinae Poey 1861 (commonly spelled Anthiinae), within the genus Tosanoides. The genus currently includes four nominal species: Tosanoides filamentosus Kamohara, 1953 (type species), T. flavofasciatus Katayama & Masuda, 1980, T. obama Pyle, Greene & Kosaki, 2016, and T. aphrodite Pinheiro, Rocha & Rocha, 2018. Herein we describe the fourth member of the genus, Tosanoides annepatrice, based on morphologic and genetic differences compared with the other known species. In addition, the species currently referred to as Pseudanthias fucinus (Randall & Ralston, 1985) from the Hawaiian Islands is likely a member of the genus Tosanoides, and is similar in many respects to T. aphrodite.
At least two other undescribed species of this genus have been reported; one from the Coral Sea (Gerald R. Allen pers. comm.) and one from Rapa Nui (Easter Island) (
Specimens were collected with hand nets during deep dives using mixed-gas, closed-circuit rebreathers, and brought to the surface alive with the aid of a hypodermic needle to vent gas from the swim bladders. Methods for counts and measurements follow those described in
The holotype has been deposited at the Bernice Pauahi Bishop Museum fish collection, Honolulu (
Fresh tissue samples were obtained from the holotype and paratypes. DNA barcodes (cytochrome c oxidase I; COI) were sequenced following the protocol described in
Republic of Palau, Kayangel State, Ngaruangl Atoll, 8.14733°N, 134.61763°E.
Holotype.
A species of Tosanoides (sensu
Dorsal fin X,16 (17), last soft ray branched to base; anal fin III,8, last soft ray branched to base; pectoral-fin rays 14; pelvic-fin rays I,5; principal branched caudal rays 14, upper procurrent unbranched caudal rays 6, lower procurrent unbranched caudal rays 4; pored lateral-line scales 33 (33–34); scale rows above lateral line to origin of dorsal fin 4; scale rows below lateral line to origin of anal fin 15 (14–15); gill rakers on upper limb 11, on lower limb 22 (22–23); vertebrae 26 (10+16).
Body ovoid, compressed, its greatest depth 2.62 (2.57–2.64) in SL, the width just posterior to gill opening 2.40 (2.22–2.56) in depth; head length 2.93 (2.35–2.77) in SL; snout short, its length 5.66 (4.28-7.00) in head; orbit diameter 2.41 (2.53–2.85) in head; interorbital convex, the least bony width 3.42 (3.45–5.41) in head; least depth of caudal peduncle 2.78 (2.99–3.40) in head.
Mouth large, oblique and protractile; lower jaw not projecting beyond the upper when mouth closed; maxilla 2.01 (1.92–2.20) in head, expanded distally, reaching below posterior border of pupil, slightly diagonal, the gape forming an angle of ca. 20° to the horizontal, supramaxilla absent. A pair of nostrils on either side of head, close together, directly in front of eye, anterior nostril in a membranous tube with an elevated posterior edge, posterior nostril with a slight rim anteriorly. Teeth in upper jaw villiform, forming a band broader anteriorly with a pair of canines on each side and another pair of canines slightly posteriorly and internally directed backward, an outer row of ca. ten slender canines on each side of jaw curved forward; lower jaw with a patch of villiform teeth anteriorly; one canine on each side anteriorly facing forward and a second canine on each side curved forward, an outer row of ca. 15 slender canines like those of the upper jaw, posterior ones pointing forward; small teeth on vomer and palatines, teeth on vomer in a triangular band; tongue pointed, smooth. Preopercle with a round angle, upper limb serrate with ca. 25 spinules, lower limb smooth; opercle with two flat spines, upper one longest and at apex; subopercle and interopercle smooth. Gill rakers long and numerous, with eleven rakers on the upper limb and 22 (22–23) on the lower limb, longest raker much longer than gill filament.
Dorsal fin very slightly notched, inserted slightly posterior to dorsal end of gill opening, its base 1.69 (1.65–1.73) in SL; first dorsal-fin spine the longest, 1.81 (1.97–2.32) in head, second dorsal-fin spine originating immediately posterior to first, its length 1.97 (2.00–2.48) in head, third dorsal-fin spine 2.13 (2.11–2.53) in head, fourth dorsal-fin spine, 2.29 (2.24–2.70) in head, fifth dorsal-fin spine 2.51 (2.33–2.76) in head, last dorsal-fin spine 2.83 (2.90–3.14) in head; membranes between dorsal-fin spines mildly incised; longest dorsal soft ray (seventh or eighth) 2.15 (2.58-2.78) in head. Anal fin originating below base of third or fourth dorsal soft ray; second anal spine slightly stronger than the third; length of first anal-fin spine 5.32 (5.17–6.00) in head, second anal-fin spine 2.13 (2.00–2.25) in head, third anal-fin spine 2.21 (2.19–2.59) in head; posterior margin of anal fin rounded; length of longest anal soft ray (fifth or sixth) 1.60 (1.77–2.38) in head. Pectoral fins subsymmetrical, longer than head, reaching a vertical at base of third or fourth anal soft ray, their length 2.31 (2.28–2.52) in SL; caudal fin deeply convex, upper and lower lobes each with two filamentous extensions on their outermost principle rays, its length 1.90 (1.95–2.26); pelvic spine 1.83 (1.66–2.15) in head; first soft ray of pelvic fin with a filamentous extension (broken in holotype), its length (3.07–4.08), in SL.
Scales moderately large, ctenoid; four in a series from origin of dorsal fin to lateral line, 15 (14–15) from origin of anal fin to lateral line; head closely scaled except for lips and tip of snout anterior to nostrils; dorsal and anal fins with small scales basally, a single row on spinous portion of dorsal fin, reaching distally ca. 1/5 of distance to outer margin posteriorly on soft portions of dorsal and anal fins; ca. seven or eight vertical scale rows on base of caudal fin; scales on pectoral fins basally, extending posteriorly on lower half of pectoral fin approximately one third distance to posterior margin. Lateral line with 33 (33–34) pored scales, high, concurrent with back, forming an angle below last several dorsal rays and extending along middle of caudal peduncle to base of caudal fin.
The two immature paratypes differ from the adult specimens in having a proportionally larger head (2.35–2.77 in SL, compared with 2.77–2.93), shorter snout (6.33–7.00 in head, compared with 4.28–5.66), longer dorsal-fin base (1.70–1.73 in SL, compared with 1.65–1.69) and anal-fin base (4.65–4.91 in SL, compared with 4.91–5.05), narrower caudal peduncle (3.17–3.40 in head, compared with 2.78–2.99), shorter pelvic spine (2.09–2.15 in head, compared with 1.66–1.83), longer pelvic fin (2.43–2.62 in head, compared with 3.07–4.08), shorter dorsal-fin rays (longest 2.77–2.78 in head, compared with 2.15–2.58) and anal-fin rays (longest 2.28–2.38 in head, compared with 1.60–1.77), shorter first anal-fin spine (5.95–6.00 in head, compared with 5.17–5.32), and shorter caudal-fin (2.08–2.26 in SL, compared with 1.90–1.95).
Color in life as in Fig.
Color in alcohol uniformly pale yellow except for eye, which is black.
Morphometric data for selected characters of type specimens are provided in Table
Morphometric and meristic data for selected characters of type specimens of Tosanoides annepatrice. Values of morphometric data (other than TL and SL) are represented as % of SL.
Measurements | Holotype | Paratype | Paratype | Paratype |
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Morphometrics | ||||
Sex | Male | Male | Immature | Immature |
Total length (TL) in mm | 80.9 | 104.0 | 46.8 | 40.4 |
Standard length (SL) in mm | 53.0 | 68.7 | 31.6 | 28.0 |
Head length | 34.2 | 36.1 | 36.1 | 42.5 |
Body depth | 38.1 | 38.7 | 38.9 | 37.9 |
Body width | 41.6 | 45.1 | 39.0 | 43.4 |
Snout length | 17.7 | 23.4 | 15.8 | 14.3 |
Predorsal length | 38.3 | 38.1 | 37.3 | 39.3 |
Preanal length | 68.1 | 63.2 | 61.7 | 65.7 |
Base of dorsal fin | 59.2 | 60.7 | 57.9 | 58.9 |
Base of anal fin | 19.8 | 20.4 | 21.5 | 20.4 |
Orbit diameter | 41.4 | 35.1 | 39.5 | 36.1 |
Interorbital width | 29.3 | 27.4 | 28.9 | 18.5 |
Caudal peduncle depth | 35.9 | 33.5 | 31.6 | 29.4 |
Pelvic spine | 54.7 | 60.1 | 46.5 | 47.9 |
Pelvic fin | 24.5 | 32.6 | 41.1 | 38.2 |
First dorsal spine length | 55.2 | 43.1 | 50.9 | 43.7 |
Second dorsal spine length | 50.8 | 43.1 | 50.0 | 40.3 |
Third dorsal spine length | 47.0 | 42.3 | 47.4 | 39.5 |
Fourth dorsal spine length | 43.6 | 37.1 | 44.7 | 38.7 |
Fifth dorsal spine length | 39.8 | 36.3 | 43.0 | 37.8 |
Last dorsal spine length | 35.4 | 31.9 | 34.2 | 34.5 |
Longest dorsal ray length | 46.4 | 38.7 | 36.0 | 36.1 |
First anal spine length | 18.8 | 19.4 | 16.7 | 16.8 |
Second anal spine length | 47.0 | 45.2 | 50.0 | 44.5 |
Third anal spine length | 45.3 | 40.7 | 45.6 | 38.7 |
Longest anal ray length | 62.4 | 56.5 | 43.9 | 42.0 |
Caudal fin length | 52.6 | 51.4 | 48.1 | 44.3 |
Pectoral fin length | 43.2 | 39.7 | 42.4 | 43.9 |
Meristics | ||||
Dorsal spines | X | X | X | X |
Dorsal rays | 16 | 17 | 17 | 17 |
Anal spines | III | III | III | III |
Anal rays | 8 | 8 | 8 | 8 |
Pectoral rays | 14 | 14 | 14 | 14 |
Pored lateral line scales | 33 | 33 | 33 | 34 |
Dorsal scale rows | 4 | 4 | 4 | 4 |
Ventral scale rows | 15 | 14 | 15 | 14 |
Gill rakers | 11+22 | 11+22 | 11+23 | 11+22 |
Tosanoides annepatrice is known on the basis of four specimens, one (the holotype) collected at a depth of 115 m in Palau, and three paratypes collected at a depth of 148 m near Pohnpei. Additional individuals have been observed at depths of ~120–150 m at Pohnpei. The species likely occurs at similar depths throughout much of Micronesia, and perhaps more broadly within the tropical western Pacific; but more exploration of habitat at appropriate depths throughout this region is necessary to determine its complete geographic range.
Tosanoides annepatrice has been observed and collected along steep limestone coral-reef drop-offs at depths from 115–150 m. The paratypes were collected along a small rocky crevice near the entrance to a cave, but other individuals have been seen in similar habitats not in association with caves. Most individuals of this species have been observed in groups consisting of one apparent male and several apparent females and juveniles.
We name this species annepatrice (a noun in apposition) in honor of Anne Patrice Greene, mother of Brian D. Greene who collected all known specimens of this new species, in recognition of the support and encouragement she has consistently provided to Brian’s exploration of the deep coral reefs of Micronesia.
The morphology of this species is consistent with the diagnosis for the genus Tosanoides as presented by Katayama & Masuda 1980. Compared with Pseudanthias Bleeker, 1871 (the only other genus it resembles), T. annepatrice shares with the other three species of Tosanoides fewer pored lateral-line scales (30–34, compared with 35–52) number of anal soft rays (8, compared with 6–7), and unbranched pectoral fin rays. Morphological comparisons with other species of Tosanoides are as follows, and only compare values for adult specimens of T. annepatrice.
Tosanoides annepatrice is more similar morphologically to T. filamentosus than to T. obama (Figure
All four species of Tosanoides can also be easily distinguished from each other on the basis of life color.
Vertebrate mtDNA barcode (cytochrome oxidase I) sequences obtained from T. annepatrice reveal genetic differences of 3.9–4.0% when compared with T. filamentosus, and 9.9–12% pairwise genetic distances when compared with the other two described species of Tosanoides (Figure
Tosanoides annepatrice is yet another example of several new fish species that have been discovered on deep coral reefs over the past several decades, mostly involving the use of modern mixed-gas closed-circuit rebreather diving technology (
We are grateful to Sonia J. Rowley, who led a series of deep-diving expeditions to Pohnpei (including the 2016 expedition on which the type specimens of T. annepatrice were collected). We are also grateful to Luiz A. Rocha for providing aquarium photos of this species, including those represented here as Figure