Research Article |
Corresponding author: Bart Shepherd ( bshepherd@calacademy.org ) Academic editor: David Morgan
© 2020 Bart Shepherd, Tyler A. Y. Phelps, Hudson T. Pinheiro, Claudia R. Rocha, Luiz A. Rocha.
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:
Shepherd B, Phelps TAY, Pinheiro HT, Rocha CR, Rocha LA (2020) Two new species of Plectranthias (Teleostei, Serranidae, Anthiadinae) from mesophotic coral ecosystems in the tropical Central Pacific. ZooKeys 941: 145-161. https://doi.org/10.3897/zookeys.941.50243
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Two new species of Plectranthias perchlets are described, collected from mesophotic coral ecosystems in French Polynesia and the Republic of the Marshall Islands, in the tropical Central Pacific. Plectranthias polygonius sp. nov. was collected at a depth of 105 m in Tahiti, French Polynesia, and 120 m in Maloelap Atoll, Republic of the Marshall Islands. It was also observed in Moorea and Rangiroa (French Polynesia), and at Majuro and Erikub Atolls, Republic of the Marshall Islands. Plectranthias hinano sp. nov. was collected at a depth of 90–98 m in Tahiti, French Polynesia, and observed in Moorea. The barcode fragment of the cytochrome oxidase I gene of Plectranthias polygonius sp. nov. does not closely match any published sequence of Plectranthias, with approximately 15% uncorrected divergence from several species. Plectranthias polygonius sp. nov. can be distinguished from all of its congeners by coloration and morphology. The barcode fragment of the COI gene of Plectranthias hinano sp. nov. is closest to Plectranthias bennetti, with 5.4% uncorrected divergence. Plectranthias hinano sp. nov. is also distinguished from all of its congeners by morphology, and a coloration that includes two indistinct black spots along the base of the dorsal-fin, and transparent yellow dorsal and anal fin membranes. With this publication, the genus Plectranthias now comprises 58 valid species, with representatives from tropical to temperate waters of the Atlantic, Pacific, and Indian oceans. These two new discoveries add to the growing body of research highlighting the rich biodiversity of mesophotic ecosystems.
biodiversity, closed-circuit rebreather, coral-reef twilight zone, ichthyology, perchlet, reef fish, taxonomy
The anthiadine genus Plectranthias Bleeker, 1873, currently comprises 56 valid species from tropical to temperate waters in the Atlantic, Pacific, and Indian oceans (
Mesophotic coral ecosystems (MCEs) are coral reef habitats found at depths of 30–150 m, and are also known as the coral reef “twilight zone” (
All specimens were collected with hand nets while diving on mixed-gas closed-circuit rebreather (Hollis Prism 2) in French Polynesia (Tahiti, Moorea) and Micronesia (Majuro, Republic of Marshall Islands). Specimens were collected and immediately transported to a field laboratory, where they were photographed, tissues sampled, fixed in 10% formalin, and preserved in 75% ethanol. The preserved specimens were later measured and x-radiographed at the California Academy of Sciences. Measurements were taken with digital calipers to the nearest 0.01 mm and rounded to one decimal place, following the conventions described in
Proportional measurements of type specimens of Plectranthias polygonius sp. nov., and Plectranthias hinano sp. nov. expressed as a percentage of the standard length.
Plectranthias polygonius sp. nov. | Plectranthias hinano sp. nov. | |||
---|---|---|---|---|
Holotype | Paratype | Holotype | Paratype | |
CAS 247193 | USNM 445722 | CAS 247195 | USNM 445723 | |
Standard length (mm) | 29.5 | 32.3 | 49.6 | 28.0 |
Head length | 31.3 | 32.3 | 42.1 | 39.8 |
Greatest body depth | 29.5 | 31.6 | 34.0 | 33.2 |
Body width | 11.1 | 13.5 | 16.5 | 13.9 |
Snout length | 9.3 | 9.0 | 14.2 | 12.3 |
Postorbital of head | 21.0 | 21.1 | 18.6 | 17.9 |
Bony interorbital width | 5.9 | 7.1 | 4.0 | 5.4 |
Orbit diameter | 11.2 | 11.5 | 11.5 | 11.4 |
Upper jaw length | 17.6 | 19.1 | 19.0 | 19.1 |
Maxilla width | 5.6 | 6.2 | 5.9 | 5.6 |
Caudal peduncle length | 8.7 | 8.1 | 10.6 | 9.7 |
Caudal peduncle depth | 12.3 | 13.0 | 11.4 | 11.8 |
Predorsal length | 38.0 | 41.4 | 41.8 | 41.0 |
Preanal length | 70.0 | 70.6 | 68.4 | 67.4 |
Prepelvic length | 35.9 | 38.7 | 42.5 | 39.9 |
Dorsal-fin base length | 48.7 | 52.9 | 48.5 | 53.7 |
First dorsal spine | 5.6 | 6.3 | 5.7 | 10.7 |
Longest dorsal spine (number) | 18.8 (3rd) | 20.0 (3rd) | 18.2 (4th) | 15.7 (3rd) |
First segmented dorsal ray | 17.1 | 15.6 | 14.9 | 16.4 |
Longest segmented dorsal ray (number) | 17.3 (2nd) | 15.6 (1st) | 15.3 (2nd) | 19.8 (3rd) |
Anal fin base length | 15.8 | 15.4 | 16.0 | 18.8 |
First anal spine | 7.6 | 8.0 | 6.3 | 8.0 |
Second anal spine | 19.0 | 19.6 | 20.6 | 20.8 |
Third anal spine | 13.2 | 13.7 | 15.9 | 15.4 |
First segmented anal ray | 17.5 | 17.7 | 16.1 | 17.7 |
Longest anal spine (number) | 25.8 (2nd) | 23.5 (2nd) | 15.4 (2nd) | 27.1 (2nd) |
Longest segmented anal ray (number) | 23.6 (4th) | 21.5 (4th) | 14.0 (2nd) | 24.8 (2nd) |
Caudal-fin length | 20.9 | 21.7 | 32.5 | 27.9 |
Pectoral-fin length | 26.2 | 29.7 | 35.7 | 38.7 |
Pelvic spine length | 16.0 | 16.5 | 17.4 | 17.4 |
Pelvic-fin length | 21.8 | 23.6 | 21.5 | 24.9 |
Mitochondrial cytochrome c oxidase subunit I (COI) DNA was sequenced and analyzed for the new species. DNA extraction and PCR amplification of the COI gene were performed following protocols detailed in
Tahiti, French Polynesia.
CAS 247193, field code: HTP906, GenBank MN922331. 29.5 mm SL, Tahiti, French Polynesia, 17°29'27"S, 149°28'01"W, depth of collection 105 m, collected with hand nets by B Shepherd, HT Pinheiro, TAY Phelps, MV Bell, and LA Rocha, 03 March 2019.
USNM 445722, field code: HTP942, GenBank MN922330. 32.3 mm SL, Maloelap Atoll, Republic of the Marshall Islands, 8°37'42"N, 170°59'58"E, depth of collection 120 m, collected with hand nets by HT Pinheiro, TAY Phelps, MV Bell, and LA Rocha, 13 August 2019.
Plectranthias polygonius sp. nov. can be distinguished from all of its congeners by live coloration, in particular the two rows of orange rhomboid-shaped polygons on the lateral part of the body and an elongated yellow and white third dorsal spine (Fig.
Proportional measurements for the type specimens are presented in Table
Body moderately elongate, laterally compressed; depth of body 3.4 in SL (3.2); width of body 2.7 in depth (2.3); head length 3.2 in SL (3.1); snout length 3.4 in HL (3.6); bony interorbital width 1.6 in snout length (1.3); orbit diameter 2.8 in HL; post-orbital head length 4.8 in SL (4.7); least depth of caudal-peduncle 2.5 in HL; caudal peduncle length 3.6 in HL (4.0).
Scales ctenoid; lateral line complete and broadly arched over pectoral fin following body contour; 30 (27) tubed scales; scales above lateral line to origin of dorsal fin 2; scales above lateral line to base of middle dorsal spine 2; scales below lateral line to origin of anal fin 9 (10); diagonal rows of scales on cheek 5; scales on top of head extending anteriorly to vertical from anterior margin of pupil; no scales on chin, maxilla, or snout; circumpeduncular scales 10 (11); gill rakers 5+13 (6+13); vertebrae 10+16; supraneurals 3; anterior supraneural-dorsal ray-pterygiophore-neural spine interdigitation pattern: 0/0+0/2/1+1/1.
Mouth large and terminal, slightly upturned; lower jaw protrudes slightly; maxilla expanded posteriorly, extending to below the posterior edge of eye; dorsal profile of head almost straight; upper jaw with one fixed, stout canine on either side of symphysis; upper canines flanked internally by villiform band with four or five rows of depressible, smaller, sharp-tipped teeth; inner rows become progressively longer, innermost row with largest teeth; lower jaw has pair of fixed, short stout canines on either side of symphysis followed by smaller, depressible, sharp-tipped conical teeth in a villiform band of 3–5 rows; lower teeth become progressively longer on inner rows; vomer roughly V-shaped band of two rows of similarly sized, sharp-tipped, conical teeth; palatines with one row of small, sharp-tipped conical teeth; tongue small, slender, pointed, and without teeth.
Opercle with three spines, the middle spine the longest; preopercle with 14 (17) small spines (serrae) along posterior margin; antrorse spines lacking on ventral margin; interopercle with no spines; subopercle smooth, with no spines; anterior nostrils positioned halfway between snout and eye, each with a small rounded flap rising from anterior rim; posterior nostrils an elliptical opening at anterior border of orbit.
Body
: overall white with two rows of bright orange rhomboid-shaped polygons, four to six in each row, arranged in an irregular grid along lateral midline of body; uppermost row of orange polygons proceeds from behind eye to dorsal third of caudal peduncle; lower row starts just dorsal to origin of pectoral fin and continues to ventral half of caudal peduncle; throat and belly white. Head: dorsal third of head orange and bottom two thirds pinkish white with two yellow stripes, both originating from the tip of the upper lip. The first extending horizontally across orbit, bifurcating past posterior edge of pupil to approximately edge of opercle. The second from tip of upper lip, tracing obliquely along maxilla and extending to ventral edge of preopercle. Preopercle region with a yellow triangular patch, from lower mid-orbit expanding in width to edge of preopercle with bifurcations to horizontal edge of operculum and pelvic fin bases respectively; orbit mostly orange-red; iris outlined in silver-grey to black with horizontal yellow stripe through middle of anterior portion; posterior portion of iris with two yellow stripes arising from a bifurcation of the anterior yellow stripe; pupil black. Fins: first three membranes of spinous portion of dorsal fin mostly orange with yellow highlights; third spine with yellow and white membrane; remaining membranes of spinous portion mostly yellow, with hyaline tips; yellow coloration continues on lower third of soft portion of dorsal fin, with upper two-thirds mostly hyaline; some yellow and pale orange on tips of soft dorsal and membranes of last four to five rays; caudal fin hyaline with white and orange rays; anal fin mostly yellow, with white margin; pelvic fins mostly white distally, with yellow rays proximally; pectoral fins hyaline. Living specimen photographed in the Marshall Islands exhibits more yellow coloration on head, within rhomboid-shaped polygons on lateral midline, and on first three membranes of spinous dorsal fin (Fig.
Light tan overall, with no visible markings.
Plectranthias polygonius sp. nov. is named for the orange rhomboid-shaped polygons arranged in parallel rows along the lateral midline that distinguish it from all other known species within the genus. To be treated as a noun in apposition.
Plectranthias polygonius sp. nov. appears to be the same species as an undescribed Plectranthias species that was photographed off Rangiroa, French Polynesia, at a depth of 65 m (
The general body shape, color, and prolongation of the third dorsal-fin spine in Plectranthias polygonius sp. nov. resemble P. inermis and P. altipinnatus (Fig.
Tahiti, French Polynesia
CAS 247195, field code: HTP909, GenBank MN922329. 49.6 mm SL, Tahiti, French Polynesia, 17°29'27"S, 149°28'01"W, depth of collection 98 m, collected with hand nets by B Shepherd, HT Pinheiro, TAY Phelps, MV Bell, and LA Rocha, 03 March 2019 (Fig.
USNM 445723, field code: TAH007, GenBank MN922328. 28.0 mm SL, Tahiti, French Polynesia, 17°36'59"S, 149°37'13"W, depth of collection 90 m, collected with hand nets by B Shepherd, HT Pinheiro, TAY Phelps, MV Bell, and LA Rocha, 28 February 2019.
Plectranthias hinano sp. nov. can be distinguished from all of its congeners by the following combination of characters: dorsal-fin rays X, 15, the last branched to base and counted as one; pectoral fin rays 11 or 12; vertebrae 10+16; lateral line complete with 29–30 tubed scales; gill rakers 7–8+12–13; ventral margin of preopercle with three antrorse spines; snout moderately long, 3.0–3.2 in HL, 12.3–14.2% SL, overall red coloration with two indistinct black spots along the base of dorsal fin, translucent yellow dorsal and anal-fin membranes.
Proportional measurements for the type specimens are presented in Table
Body moderately elongate, laterally compressed; depth of body 2.9 in SL (3.0); width of body 2.1 in depth (2.4); head length 2.4 in SL (2.5); snout length 3.0 in HL (3.2); bony interorbital width 3.5 in snout length (2.3); orbit diameter 3.7 in HL (3.5); post-orbital head length 5.4 in SL (5.6); least depth of caudal-peduncle 3.7 in HL (3.4); caudal-peduncle length 4.0 in SL (4.1).
Scales ctenoid; lateral line complete and broadly arched over pectoral fin following body contour; 29 (30) tubed scales; scales above lateral line to origin of dorsal fin 3; scales above lateral line to base of middle dorsal spine 2; scales below lateral line to origin of anal fin 12; diagonal rows of scales on cheek 4; scales on top of head extending anteriorly to vertical from center of eye; area on top of head between eyes with scales; no scales on chin, maxilla, or snout; circumpeduncular scales 12; gill rakers 8+13 (7+12); vertebrae 10+16; supraneurals 3; anterior supraneural-dorsal ray-pterygiophore-neural spine interdigitation pattern: 0/0+0/2/1+1/1.
Mouth large and terminal, slightly upturned; lower jaw protrudes slightly; maxilla expanded posteriorly, extending to below the posterior edge of pupil; upper jaw with one fixed, stout canine on either side of symphysis; upper canines flanked internally by villiform band with 7–9 irregular rows of depressible, smaller, sharp-tipped teeth; inner rows become progressively longer, innermost row with largest teeth; lower jaw has one large, fixed canine on either side of lower jaw, roughly at midpoint, followed by smaller, depressible, sharp-tipped conical teeth in a villiform band of 4–6 irregular rows, innermost teeth same size as outer rows; vomer roughly V-shaped band of two rows of similarly-sized, sharp-tipped, conical teeth; palatines with one row of small, sharp-tipped conical teeth anteriorly, two rows posteriorly; tongue small, slender, pointed, and without teeth.
Opercle with three flat spines, the middle spine the longest; preopercle with 22 (12) small spines (serrae) along posterior margin; ventral (inferior) margin of preopercle with three antrorse spines; interopercle with no spines; subopercle smooth, with one spine on inferior margin; anterior nostrils positioned closer to eye than to snout, each with a small rounded flap rising from anterior rim; posterior nostrils an elliptical opening at anterior border of orbit.
Body
: overall light red in color; chest and belly mostly light red to pink; dorsal portion of body darker red with yellow along scale margins and lateral line; series of 8–12 pink to white incomplete bars on the body, originating just behind orbit and continuing to base of caudal fin; bars are approximately 20 degrees off of vertical anteriorly, becoming near vertical as they approach base of caudal fin; black spot, almost twice the diameter of orbit, at base of spinous dorsal-fin spines 7–10 and soft rays 1–5, continuing slightly more than halfway up membrane of spinous dorsal; black spot is interrupted by the near-vertical white bar originating below soft rays 1 or 2; smaller second black spot, slightly smaller than orbit, located at base of soft dorsal rays 11–15. Head: snout, throat, anterior portion of lower lip, maxilla, and operculum mostly light pink; yellow stripe originating at snout, proceeding across maxilla, below orbit to preopercle; lattice-like network of indistinct yellow stripes radiating outward from pupil across iris, between eyes, across top of head, from ventral margin of orbit to origin of lateral line and lower margin of operculum; iris mostly pink with yellow splotches radiating outward from pupil; pupil black and teardrop-shaped, pointed anteriorly. Fins: spinous portion of dorsal fin predominantly translucent yellow, with upper half of black spot on membrane between spines 7–10; lower third of soft dorsal fin mostly translucent yellow, upper two-thirds hyaline; caudal-fin membranes mostly hyaline with some regions a faint transparent yellow, fin rays white with red margins; pelvic fins hyaline with faint yellow on rays; anal fin mostly translucent yellow with hyaline margins; pectoral fins hyaline with rays outlined in pink; base of pectoral fins yellow. Freshly dead specimens exhibit similar coloration, with slightly more yellow on head, body, and fins (Fig.
Light tan overall, with dark brown stippling in vertical bands along lateral sides of body, darkest brown along base of dorsal fin at location of black spots.
Plectranthias hinano sp. nov. is named after Teurumereariki Hinano Teavai Murphy, former associate director of the University of California Berkeley Gump Research Station and president of the cultural association Te Pu Atitia, in honor of the significant contributions she has made supporting Polynesian biocultural heritage and field research in Moorea, French Polynesia. The name is a noun in the genitive case.
The two specimens described in this paper, plus an individual that was retained for public aquarium exhibition, were collected respectively in Tahiti and Moorea, French Polynesia. A similar species was photographed at 120 m depth in Erikub Atoll, Republic of Marshall Islands, however specimens were not retained (Fig.
Living specimens of A Plectranthias hinano sp. nov. at Steinhart Aquarium, California Academy of Sciences B Plectranthias cf. bennetti photographed at a depth of 120 m at Maloelap Atoll, Republic of the Marshall Islands, and C Plectranthias bennetti photographed in an aquarium. Photographs by T Wong (A), LA Rocha (B), and YK Tea (C).
The most similar barcode fragment of the mtDNA COI gene to Plectranthias hinano sp. nov. is P. bennetti from the Coral Sea (5.5% uncorrected pairwise genetic distance). Morphologically, it can be distinguished from P. bennetti by having a longer snout (3.0–3.2 in HL vs. 4.4 in P. bennetti), a larger orbit (3.5–3.7 in HL vs. 4.1 in P. bennetti), in the number of circumpeduncular scales (12 vs. 14 in P. bennetti), the number of gill rakers (7–8 + 12–13, vs. 5+13 in P. bennetti), and in coloration (by having two indistinct black spots along the base of the dorsal fin, and yellow dorsal and anal fin membranes).
Species within the genus Plectranthias appear to be common and conspicuous inhabitants of MCEs, and we have regularly observed them in highly complex, rocky habitats with abundant coralline algae, sponges, and black corals (Fig.
More than 70% of all research on MCEs has been published in the past seven years (
Plectranthias inermis. CAS 241326. Philippines, Batangas, 28 April 2015.
P. japonicus. CAS 33555, 1. Philippines, Batangas, 23 June 1966.
P. longimanus. CAS 213185, 3. Fiji, Viti Levu Island, 31 May 1999.
P. sagamiensis. CAS 235596, 1. Philippines, Luzon Island, 02 June 2011.
P. winniensis. CAS 219169, 1. Fiji, Vanua Levu Island, 27 May 2003.
This work was funded by the generous support of donors to the California Academy of Sciences’ Hope for Reefs Initiative. Our research in French Polynesia was conducted under permit number USR 3278 from the Ministry of Fisheries, and in the Republic of the Marshall Islands under permit from the Marshall Islands Marine Resources Authority. We are grateful to many colleagues who helped in the field, lab, and with discussions: MV Bell, M Bozinovic, C Castillo, D Catania, K de Brum, J Fong, E Kabua-Tibon, M Lane, A Mecho, JE McCosker, A Shafer, G Siu, B Stockwell, YK Tea, H Teavai-Murphy, M White, N Yim, and S Yong. Logistics, equipment, and technical diving support was provided by Hollis Rebreathers LLC, G Siu, L Roattino, C Cicculo, SubTecnologie, and Indies Trader. We would like to thank J Fong for taking the radiographs, A Gaisiner for photographing the preserved types, T Sinclair-Taylor for photographing the Marshall Islands specimen and field support, T Wong for photographing the living specimen at Steinhart Aquarium, and YK Tea for providing photographs of closely related species and valuable suggestions to improve the manuscript.