Research Article |
Corresponding author: Dongsheng Zhang ( dszhang@sio.org.cn ) Academic editor: Sabine Stöhr
© 2022 Xiaojun Xie, Bo Lu, Jie Pang, Dongsheng Zhang.
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:
Xie X, Lu B, Pang J, Zhang D (2022) Description of two species of the genus Astrodia Verrill, 1899 (Ophiuroidea, Euryalida, Asteronychidae), including a new species from seamounts in the West Pacific. ZooKeys 1123: 99-122. https://doi.org/10.3897/zookeys.1123.87397
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Five specimens of Ophiuroidea from deep-sea seamounts in the West Pacific were collected and identified as two species, Astrodia duospina sp. nov. and Astrodia abyssicola. The new species, Astrodia duospina sp. nov., can be distinguished from its congeners by having indistinct or underdeveloped oral papillae, relatively short genital slits, crescent-shaped lateral arm plates, and plate-shaped external ossicles on the aboral surface of the disc. One specimen was identified as Astrodia abyssicola, which has been reported in the north-western Pacific and the north-eastern coast of Japan. The most recent tabular key of Astrodia was revised with two more key characteristics added, the shape and presence of oral papillae and the number of arm spines. The phylogenetic relationship of Astrodia and Asteronyx was analyzed based on 16S and COI sequences. The discovery of the two species further expanded the geographical distribution of the genus Astrodia.
Deep sea, molecular phylogeny, morphology, ophiuroids, taxonomy
Class Ophiuroidea, as the largest group among echinoderms, with 2126 valid species (
The order Euryalida Lamarck, 1816 comprises about 200 species from three families, Euryalidae Gray, 1840, Asteronychidae Ljungman, 1867, and Gorgonocephalidae Ljungman, 1867 (
In this study, we describe a new species, Astrodia duospina sp. nov., and redescribe Astrodia abyssicola, from seamounts of the West Pacific. New interspecific diagnostic characteristics were identified, and the tabular key of
Five specimens of Astrodia were collected by ROV HAILONG III, ROV HAILONG IV, and HOV JIAOLONG, from seamounts in the Philippine Sea and the Northwest Pacific, during several COMRA’s cruises in 2013, 2020, and 2021 (Fig.
Morphological characters were examined and photographed using a stereoscopic microscope (Zeiss Axio Zoom V16). Arm skeletons were examined with a Hitachi TM1000 scanning electron microscope. Skeletal elements were prepared by submerging in commercial bleach (2.5% NaOCl). Washed in distilled water and ethanol, air-dried, and mounted on a stub using dissolved carbon tapes. The following literature was used as references for the morphological analysis:
Several arm segments were dissected from each individual for genomic DNA extraction using DNeasy Blood & Tissue Kit (QIAGEN) following the manufacturer’s protocols. The COI sequences and 16S rRNA sequences were amplified with primers listed in Table
Primer | Sequence |
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Oph-COI-F | TTTCAACTAATCAYAAGGAYATWGG |
Oph-COI-R | CTTCAGGRTGWCCRAARAAYCA |
16Sar | CGCCTGTTTATCAAAAACAT |
16Sbr | CCGGTCTGAACTCAGATCACGT |
Seventy-two 16S sequences and 28 COI sequences of Asteronychidae were downloaded from the NCBI. In total, 78 16S sequences and 34 COI sequences (Table
Voucher specimens and accession numbers of COI and 16S sequence data used in the phylogenetic analysis (IDSSE, Institute of Deep-sea Science and Engineering, China; MV, Museums Victoria, Australia; NSMT, National Museum of Nature and Science, Japan; RSIO, Second Institute of Oceanology, China;
Species | Locality | Voucher number | CO1 | 16S | Code from |
---|---|---|---|---|---|
Asteronyx longifissus | Monterey, California |
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- | KM014337 | - |
Asteronyx loveni | South China Sea | IDSSE-EEB-SW0002 | MZ198756 | MZ203264 | - |
Asteronyx loveni | New Zealand | MV F188855 | KU895061 | - | - |
Asteronyx loveni | Off Abashiri, Hokkaido | NSMT E-6904-A | - | LC276316 | OK-226 |
Asteronyx loveni | Off Abashiri, Hokkaido | NSMT E-6904-B | - | LC276354 | OK-315 |
Asteronyx loveni | Off Abashiri, Hokkaido | NSMT E-6904-C | LC276289 | LC276330 | OK-256 |
Asteronyx loveni | Off Abashiri, Hokkaido | NSMT E-6904-G | LC276290 | LC276331 | OK-257 |
Asteronyx loveni | Off Abashiri, Hokkaido | NSMT E-6904-H | LC276282 | LC276317 | OK-227 |
Asteronyx loveni | Off Abashiri, Hokkaido | NSMT E-6904-I | - | LC276359 | OK-339 |
Asteronyx loveni | Off Abashiri, Hokkaido | NSMT E-6904-J | - | LC276350 | OK-295 |
Asteronyx loveni | Off Abashiri, Hokkaido | NSMT E-6904-K | - | LC276332 | OK-258 |
Asteronyx loveni | Off Abashiri, Hokkaido | NSMT E-6904-L | - | LC276358 | OK-337 |
Asteronyx loveni | Off Abashiri, Hokkaido | NSMT E-6904-R | - | LC276334 | OK-262 |
Asteronyx loveni | Off Abashiri, Hokkaido | NSMT E-6904-S | - | LC276353 | OK-314 |
Asteronyx loveni | Off Abashiri, Hokkaido | NSMT E-6904-T | - | LC276333 | OK-261 |
Asteronyx loveni | Off Abashiri, Hokkaido | NSMT E-6951-B | - | LC276343 | OK-281 |
Asteronyx loveni | Off Abashiri, Hokkaido | NSMT E-6951-C | LC276292 | LC276337 | OK-269 |
Asteronyx loveni | Off Abashiri, Hokkaido | NSMT E-6951-D | - | LC276344 | OK-284 |
Asteronyx loveni | Off Abashiri, Hokkaido | NSMT E-6951-F | - | LC276336 | OK-268 |
Asteronyx loveni | Off Abashiri, Hokkaido | NSMT E-6951-G | - | LC276341 | OK-279 |
Asteronyx loveni | Off Abashiri, Hokkaido | NSMT E-6951-H | LC276291 | LC276335 | OK-267 |
Asteronyx loveni | Off Miyako, Iwate | NSMT E-6943-A | LC276288 | LC276329 | PT-253 |
Asteronyx loveni | Off Miyako, Iwate | NSMT E-6256 | AB758757 | AB605076 | PT-41 |
Asteronyx loveni | Off Miyako, Iwate | NSMT E-5641-A | LC276284 | LC276320 | PT-238 |
Asteronyx loveni | Off Miyako, Iwate | NSMT E-5641-B | LC276285 | LC276321 | PT-239 |
Asteronyx loveni | Off Miyako, Iwate | NSMT E-5641-C | - | LC276322 | PT-240 |
Asteronyx loveni | Off Miyako, Iwate | NSMT E-5641-D | LC276286 | LC276323 | PT-241 |
Asteronyx loveni | Off Miyako, Iwate | NSMT E-5641-E | - | LC276324 | PT-242 |
Asteronyx loveni | Off Miyako, Iwate | NSMT E-5638-A | LC276278 | LC276308 | PT-213 |
Asteronyx loveni | Off Miyako, Iwate | NSMT E-5638-B | - | LC276352 | PT-306 |
Asteronyx loveni | Off Miyako, Iwate | NSMT E-5638-D | - | LC276357 | PT-323 |
Asteronyx loveni | Off Miyako, Iwate | NSMT E-5638-E | - | LC276356 | PT-320 |
Asteronyx loveni | Off Miyako, Iwate | NSMT E-5637-A | - | LC276310 | PT-215 |
Asteronyx loveni | Off Miyako, Iwate | NSMT E-5637-B | LC276281 | LC276314 | PT-220 |
Asteronyx loveni | Shima Spur, Mie | NSMT E-6360 | - | LC276302 | PM-199 |
Asteronyx loveni | Shima Spur, Mie | NSMT E-6983 | LC276280 | LC276312 | PM-218 |
Asteronyx loveni | Shima Spur, Mie | NSMT E-6983 | - | LC276347 | PM-290 |
Asteronyx loveni | Shima Spur, Mie | NSMT E-6982 | - | LC276309 | PM-214 |
Asteronyx loveni | Off Tosa, Kochi | NSMT E-1143-A | - | LC276318 | PK-231 |
Asteronyx loveni | East China Sea, west of Japan | NSMT E-6986-A | LC276273 | LC276298 | ECS-195 |
Asteronyx loveni | East China Sea, west of Japan | NSMT E-6986-C | LC276272 | LC276297 | ECS-194 |
Asteronyx reticulata | East of Hiraji Bank, Nagasaki | NSMT E-6912 | - | LC276355 | ECS-316 |
Asteronyx reticulata | East of Hiraji Bank, Nagasaki | NSMT E-6915 | - | LC276338 | ECS-272 |
Asteronyx reticulata | East of Hiraji Bank, Nagasaki | NSMT E-7016 | - | LC276301 | ECS-198 |
Asteronyx reticulata | East of Naka-Kasayama Bank, Nagasaki | NSMT E-6908-C | - | LC276293 | ECS-190 |
Asteronyx reticulata | East of Naka-Kasayama Bank, Nagasaki | NSMT E-6908-D | LC276271 | LC276296 | ECS-193 |
Asteronyx reticulata | East of Naka-Kasayama Bank | NSMT E-6931 | LC276279 | LC276311 | ECS-217 |
Asteronyx reticulata | West of Gajajima Isl. Kagoshima | NSMT E-6354 | - | LC276305 | ECS-204 |
Asteronyx reticulata | East of Hiraji Bank, Nagasaki | NSMT E-6910 | - | LC276300 | ECS-197 |
Asteronyx reticulata | East of Hiraji Bank, Nagasaki | NSMT E-6911 | - | LC276294 | ECS-191 |
Asteronyx reticulata | East of Hiraji Bank, Kagoshima | NSMT E-6926 | - | LC276342 | ECS-280 |
Asteronyx reticulata | East of Hiraji Bank, Kagoshima | NSMT E-6929 | - | LC276304 | ECS-203 |
Asteronyx reticulata | West off Takarajima Isl. | NSMT E-6355 | LC276274 | LC276299 | ECS-196 |
Asteronyx reticulata | West of Amami Ohshima Isl., Kagoshima | NSMT E-6351-A | - | LC276325 | ECS-243 |
Asteronyx reticulata | West of Amami Ohshima Isl., Kagoshima | NSMT E-6942-B | - | LC276339 | ECS-274 |
Asteronyx reticulata | West of Ensei Knoll, Kagoshima | NSMT E-6921 | - | LC276349 | ECS-294 |
Asteronyx reticulata | West of Ensei Knoll, Kagoshima | NSMT E-6922-A | LC276287 | LC276328 | ECS-249 |
Asteronyx reticulata | West of Ensei Knoll, Kagoshima | NSMT E-6925-A | - | LC276326 | ECS-247 |
Asteronyx reticulata | West of Ensei Knoll, Kagoshima | NSMT E-6925-B | - | LC276327 | ECS-248 |
Asteronyx reticulata | East China Sea, west of Japan | NSMT E-7001 | LC276276 | LC276306 | ECS-205 |
Asteronyx reticulata | East China Sea, west of Japan | NSMT E-7002 | LC276277 | LC276307 | ECS-206 |
Asteronyx reticulata | Off Amami Ohshima Isl. Kagoshima | NSMT E-6352 | - | LC276315 | ECS-223 |
Asteronyx reticulata | West of Minami-Ensei Knoll, Kagoshima | NSMT E-6916 | - | LC276345 | ECS-286 |
Asteronyx reticulata | West of Minami-Ensei Knoll, Kagoshima | NSMT E-6923-A | - | LC276340 | ECS-278 |
Asteronyx reticulata | West of Minami-Ensei Knoll, Kagoshima | NSMT E-6923-B | LC276275 | LC276303 | ECS-202 |
Asteronyx reticulata | East China Sea, west of Japan | NSMT E-7003-A | - | LC276351 | ECS-303 |
Asteronyx reticulata | East China Sea, west of Japan | NSMT E-7003-B | - | LC276346 | ECS-288 |
Asteronyx reticulata | East China Sea, west of Japan | NSMT E-7000-A | - | LC276348 | ECS-291 |
Asteronyx reticulata | West of Minami-Ensei Knoll, Kagoshima | NSMT E-6920 | LC276270 | LC276295 | ECS-192 |
Asteronyx reticulata | Off Iejima Isl., Okinawa | NSMT E-6987 | - | LC276313 | ECS-219 |
Asteronyx sp. | Between Yakushima Isl and Tanegashima Isl., Kagoshima | NSMT E-3157-B | LC276283 | LC276319 | PSW-237 |
Asteronyx luzonicus | South China Sea | IDSSE-EEB-SW0003 | MZ198757 | MZ203265 | - |
Astrodia abyssicola | Miyagi, off Onahama | NSMT E-6257 | AB758828 | AB605077 | - |
Astrodia abyssicola | Philippine Sea, KPR Seamount | RSIO68002 | OP328783 | OP325293 | - |
Astrodia duospina sp. nov. | Philippine Sea, KPR Seamount | RSIO59012 | OP328780 | OP325290 | - |
Astrodia duospina sp. nov. | Northwest Pacific, Ko-Hakucho-Guyout Seamount | RSIO61068 | OP328781 | OP325291 | - |
Astrodia duospina sp. nov. | Northwest Pacific, RB Seamount | RSIO61069 | OP328782 | OP325292 | - |
Asteroschema ajax | Off Lord Howe Isl. | MV F99759 | AB758762 | AB605078 | - |
Asteroschema clavigerum | North Atlantic | haplotype 1 | HM587850 | HM587828 | - |
Class Ophiuroidea Gray, 1840
Order Euryalida Lamarck, 1816
Family Asteronychidae Ljungman, 1867
Holotype : China • 1 specimen; Northwest Pacific, Nazimov Guyot; 15°11.34'N, 162°49.26'E; depth 2713 m; 16 September 2020; collected by ROV HAILONG III; preserved in alcohol; RSIO61068. Paratypes: China • 1 specimen; Northwest Pacific, Nazimov Guyot; 15°11.34'N, 162°49.26'E; depth 2713 m; 16 September 2020; collected by ROV HAILONG III; preserved in alcohol; RSIO61069 • 1 specimen; Northwest Pacific, Caiwei Guyot; 15°40.61'N, 154°53.77'E; depth 2744 m; 7 September 2013; collected by HOV JIAOLONG; preserved in alcohol; RSIO31004 • 1 specimen; the Philippine Sea, Kyushu-Palau Ridge, Roischesar Peak; 13°20.85'N, 134°32.81'E; depth 1900–2000 m; 2 August 2020; collected by ROV HAILONG IV; preserved in alcohol; RSIO59012.
Disc raised high above the arm. Aboral disc with plate-shaped external ossicles in the center and on the periphery. Radial shield narrow, longer than wide. Teeth triangular, oral papillae indistinct or underdeveloped. Genital slits short, approximately one-fourth of the height of the disc. Lateral arm plates crescent and not projecting on arms. Arm spines no more than two.
Disc pentagonal, notched interradial edges, 14 mm in diameter, 4.7 mm in height. Aboral surface almost flat, slightly depressed in the center, entirely covered by thickened skin with plate-shaped external ossicles in the center, about 220 μm long (Fig.
In situ and on-board photos of Astrodia duospina sp. nov. A photo in situ (RSIO61068: the individual below, RSIO61069: the individual above, attached to an unidentified sea pen species) B photo on board (RSIO61068: the individual on the left, RSIO61069: the individual on the right) C, D photos on board (RSIO31004), aboral side (C), oral side (D). Scale bars: 10 mm (B); 20 mm (C, D).
Morphological characters of Astrodia duospina sp. nov. (holotype: RSIO61068) A aboral view of the disc B periphery of the aboral disc C oral view of the disc D genital silts E aboral view of the arms F arms spines. Abbreviations: RS radial shield; PO plate-shaped ossicle; M madreporite; T teeth; OT oral tentacle; GS genital slit; G gonad; LAP lateral arm plate; AS arm spine. Scale bars: 2 mm (A, C, E); 1 mm (B, D); 0.5 mm (F).
Oral surface flat, covered by thickened skin. Oral shield small to invisible, one madreporite. Adoral shield obscured by skin (Fig.
Five arms, long and slender, about eight to nine times as long as disc diameter, no abrupt change in width basally (Fig.
Color. Pink in situ, white in alcohol (Fig.
Ossicle morphology of holotype. Vertebrae articulation streptospondylous, wider than long in proximal segments (Fig.
Lateral arm plates and arm spines of Astrodia duospina sp. nov. (holotype: RSIO61068) A, B lateral arm plates from proximal arm, outer view (A), inner view (B) C, D lateral arm plates from distal arm, outer view (C), inner view (D) E, F arm spines from proximal (E) and distal arm (F). Abbreviations: MO muscle opening; NO nerve opening; R ridge. Scale bars: 200 μm (E); 100 μm (F); 90 μm (A, B); 60 μm (C, D).
Two paratypes (RSIO31004, RSIO61069) share the same morphological characteristics as the holotype, disc diameter 10.17 and 13.94 mm, about 1/10 and 1/9 as wide as the length of the arms, respectively. However, the radial shields of RSIO31004 are shorter than the radial shields of the holotype and of
RSIO61069 (Fig.
Morphological characters of paratypes of Astrodia duospina sp. nov. A arm spines of RSIO31004 B arm spines of RSIO61069 C–E pictures of RSIO59012, oral disc (C), aboral disc (D), arm and arm spines (E) F aboral disc of RSIO31004. These specimens have exceptionally three arm spines for an arm segment (shown by an arc in A and B). Abbreviations: AS arm spine; T teeth; PO plate-shaped ossicle; GS genital slit; G gonad; RS radial shield. Scale bars: 2 mm (F); 1 mm (C, E); 0.5 mm (A, B, D).
The species name duo is derived from the Latin numeral word, meaning two, and Latin feminine noun, spina, meaning spine, referring to the presence of no more than two arm spines throughout the arm.
This new species falls within the genus Astrodia by only possessing cylindrical unhooked arm spines. The new species resembles Astrodia abyssicola mostly by having plate-shaped external ossicles on the aboral disc and crescent-shaped lateral arm plates. However, the oral papillae are indistinct or underdeveloped in Astrodia duospina, which can be used to distinguish the two species from each other (Fig.
Astrodia tenuispina is a widely distributed species and was characterized by having slender unhooked arm spines, small and short oral papillae, separated genital slits (
Ophiocreas abyssicola Lyman, 1879: 64–65, plate 17, figs 470–473.
Astrodia abyssicola:
China • 1 specimen; Philippine Sea, Kyushu-Palau Ridge, Mugiboshi Seamount; 16.57.14'N, 134.52.7'E; depth 3225 m; 11 August 2021; collected by an HOV JIAOLONG; preserved in alcohol; RSIO68002.
Disc pentagonal and almost flat, 10 mm in diameter, 3.2 mm in height, skin wrinkled under dry conditions (Fig.
Oral surface flat, covered by thin skin, and lacking external ossicles (Fig.
Five arms, long and slender, about nine to ten times as long as disc diameter, no abrupt change in width basally (Fig.
Color. Bright pink in situ, entirely white in alcohol (Fig.
Morphological characters of Astrodia abyssicola (RSIO68002) A aboral view of the disc B center of the aboral disc C oral view of the disc D genital silts E aboral view of the arms F arms spines. Abbreviations: RS radial shield; M madreporite; T teeth; ADS adoral shield; OT oral tentacle; GS genital slit; G gonad; LAP lateral arm plate; AS arm spine. Scale bars: 2 mm (A, C, E); 1 mm (B, D, F).
Ossicle morphology. Vertebrae articulation streptospondylous, wider than long in proximal segments (Fig.
Lateral arm plates and arm spines of Astrodia abyssicola (RSIO68002) A, B lateral arm plates from the proximal arm, outer view (A), and inner view (B) C, D lateral arm plates from the distal arm, outer view (C), and inner view (D) E, F arm spines from proximal (E) and distal (F). Abbreviations: MO muscle opening; NO nerve opening; R ridge. Scale bars: 90 μm (A, B, F); 60 μm (C, D, E).
Ophiocreas abyssicola was first described by
The key morphological characters among the five species from the genus Astrodia based on
Comparison of key morphological characters among species in the genus Astrodia.
Species | Arm spines | Genital slits | External ossicles | Lateral arm plates on middle to distal portion of arms | Oral papillae | Reference |
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Astrodia abyssicola (Lyman, 1879) | 0–2 | very short, ~1/5 (height of disc) | plate-shaped on periphery | shapes: crescent; projections: absent | domed granule-shaped |
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Astrodia excavata (Lütken & Mortensen, 1899) | 0–3 | large, ~2/3 (height of disc) | granule-shaped near radial shields and genital slits | shapes: bar-like; projections: present | domed granule-shaped |
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Astrodia plana (Lütken & Mortensen, 1899) | 0–3 | short, ~1/4 (height of disc) | absent | shapes: oblong; projections: absent | domed granule-shaped |
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Astrodia tenuispina (Verrill, 1884) | 0–3, occasionally 4 | short, ~1/2 (height of disc) | plate-shaped on periphery, granule-shaped in center | shapes: unknown; projections: absent | domed granule-shaped, small and short |
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Astrodia duospina sp. nov. | 0–2, occasionally 3 | short, ~1/4 (height of disc) | plate-shaped on periphery and in center | shapes: crescent; projections: absent | indistinct or underdeveloped | This study |
Based on the COI (583~1511 bp) and 16S (431~539 bp) sequences, the phylogenetic relationship of the two genera, Astrodia and Asteronyx, was inferred. The ML tree based on the concatenated 16S and COI sequences suggested that both Astrodia and Asteronyx were monophyletic with high bootstrap values (Fig.
Maximum likelihood tree of the genus Astrodia based on concatenated sequences of COI and 16S (clades of Lin 1A, Lin 1B, Lin 2A, Lin 2B, Lin 2C and Lin 3 are from
In this study, we described a new species of the genus Astrodia collected from seamounts in the West Pacific, and another species (Astrodia abyssicola) was redescribed. Through comparing the five species of Astrodia, the tabular key of
We thank all the scientists and crew on the RVs DAYANGYIHAO and XIANGYANGHONG, and the HAILONG and JIAOLONG team for their great work in the collection of the specimens. We also like to thank Dr Timothy D. O’Hara and Dr Masanori Okanishi and Ruiyan Zhang for their precious advice and kind help. This study was supported by the National Natural Science Foundation of China (42076135), the foundation of China Ocean Mineral Resources R & D Association (No. DY135-E2-2-06).
Figure S1
Data type: Image.
Explanation note: Maximum likelihood tree of the genus Astrodia and Asteronyx based on concatenated sequences of COI and 16S.
Figure S2
Data type: Image.
Explanation note: Maximum likelihood tree of the genus Astrodia and Asteronyx based on COI sequences.
Figure S3
Data type: Image.
Explanation note: Maximum likelihood tree of the genus Astrodia and Asteronyx based on 16S sequences.