Nethupul H et al. (2022), Order Euryalida (Echinodermata, Ophiuroidea), ...

ECB-ART-50324

Zookeys 2022 Mar 30;1090:161-216. doi: 10.3897/zookeys.1090.76292.

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Order Euryalida (Echinodermata, Ophiuroidea), new species and new records from the South China Sea and the Northwest Pacific seamounts.

Nethupul H , Stöhr S , Zhang H .

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Ophiuroids were collected by the manned submersible 'Shenhaiyongshi' from the deep-sea seamounts in the South China Sea and Northwest Pacific regions at 602-1920 m depth, during 2018 to 2020. A total of nine species was identified, including two new species and seven new records from the South China Sea and one new record from the Northwest Pacific region. Two new species are described as Asteroschemashenhaiyongshii sp. nov. and Asteroschemadomogranulatum sp. nov. The seven new records included five species from the genus Asteroschema, and one species each from the genera Asterostegus and Astrodendrum. Comprehensive descriptions of morphological features are provided, including characteristics of the arm skeleton, as well as a phylogenetic analysis based on 16S and COI sequences. Intraspecific genetic distance ranges of Euryalida species from the present study were 0.34% to 1.38%, which was relatively low compared to other orders in the class Ophiuroidea. The present study suggests a high probability that species of the order Euryalida are more widely spread around the Indo-Pacific region than previously expected.

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	Figure 1. Collecting stations in this study A, A1 South China Sea (Hainan, Xisha, and Zhongsha Islands) B, B1 Northwest Pacific (southwest of Guam). Source: International Hydrographic Organization and Sieger (2012).
	Figure 2. Family Euryalidae, maximum likelihood (ML) tree based on partial COI and 16S sequences (bootstrap support values were generated with rapid bootstrapping algorithm for 1,000 replicates; blue = new species; brown = specimens from this study).
	Figure 3. Family Gorgonocephalidae, maximum likelihood (ML) tree based on partial COI sequences (bootstrap support values were generated with rapid bootstrapping algorithm for 1,000 replicates; green = specimen from this study).
	Figure 4. Asteroschemadomogranulatum sp. nov., holotype (IDSSE-EEB-SW0089) A dorsal view B ventral view C dorsal disc D lateral disc E ventral disc F dorsal arm (proximal) G lateral arm (proximal) H lateral arm (distal) I ventral arm (base) J, K ventral arm (proximal) L ventral arm (middle) M, N arm spines (middle) O, P arm spines (distal). Abbreviations: ars arm spine, arsb arm spine base, as adoral shield, ass adoral shield spine, gs genital slit, iars inner arm spine, j jaw, oars outer arm spine, po plate-like ossicle, rs radial shields, t teeth, tp tentacle pore. Scale bars: 2 mm (A, B); 1 mm (CâG, K, L); 500 Âµm (HâJ, M, N); 200 Âµm (O, P).
	Figure 5. Asteroschemadomogranulatum sp. nov., paratype (IDSSE-EEB-SW0090) A, B lateral arm plate (external, internal) C arm spines (middle) D arm spine (distal) E skin from dorsal arm base, insert frame shows polygonal plate-like large ossicle FâJ vertebrae F proximal view G distal view H lateral view I dorsal view J ventral view. Abbreviations: d dorsal, de depression, dist distal, iars inner arm spine, lac lateral ambulacral canals, mo muscle opening, no nerve opening, oars outer arm spine, pb podial basin, prox proximal, st secondary teeth, v ventral. Scale bars: 800 Âµm (FâJ); 500 Âµm (C); 300 Âµm (B, D, E); 200 Âµm (A).
	Figure 6. Asteroschemashenhaiyongshii sp. nov., holotype (IDSSE-EEB-SW0086) A dorsal view B ventral view C dorsal disc D lateral disc E oral frame F dorsal arm (base) G lateral arm (proximal) H dorsal arm (distal) I ventral arm (base) J ventral arm (middle) K ventral arm (distal) L arm spine (arm base) M, N arm spines (middle) O, P arm spines (distal). Abbreviations: ars arm spine, go granular ossicle, goc granular ossicles coat, gs genital slit, iars inner arm spine, j jaw, oars outer arm spine, rs radial shield, t teeth, tp tentacle pore. Scale bars: 2 mm (A, B); 1 mm (C, D, F, I, J); 500 Âµm (E, G, H, K, M, N, P); 200 Âµm (L, O).
	Figure 7. Asteroschemashenhaiyongshii sp. nov., paratype (IDSSE-EEB-SW0087) AâC lateral arm plate (external, internal) D, E arm spines (middle) F, G arm spine (distal) H skin from dorsal arm base (rounded granular ossicles) IâM vertebrae I proximal view J distal view K lateral view L dorsal view M ventral view. Abbreviations: d dorsal, de depression, dist distal, lac lateral ambulacral canals, mo muscle opening, no nerve opening, pb podial basin, prox proximal, st secondary teeth, v ventral. Scale bars: 500 Âµm (D, IâM); 300 Âµm (AâC, E, G); 200 Âµm (F, H).
	Figure 8. Asteroschemacf.bidwillae McKnight, 2000 (IDSSE-EEB-SW0105) A dorsal side of the specimen B ventral side of the specimen C dorsal disc D skin on the disc E lateral disc F oral frame G dorsal arms (proximal) H dorsal arm (middle) I ventral arm (arm base) J outer arm spine (proximal) K arm spines (middle) L arm spines (distal). Abbreviations: ars arm spine, as adoral shield, ass adoral shield spine, go granular ossicle, gs genital slit, iars inner arm spine, j jaw, oars outer arm spine, rs radial shield, t teeth, tp tentacle pore. Scale bars: 2 mm (AâC, E, G); 1 mm (F, H, I); 500 Âµm (K, L); 200 Âµm (D, G, J).
	Figure 9. Asteroschemacf.bidwillae McKnight, 2000 (IDSSE-EEB-SW0105) A, B lateral arm plate (external, internal) C outer arm spine (middle) D arm spine (distal) E skin from dorsal arm base (rounded to irregular-shaped granular ossicles) FâJ vertebrae F proximal view G distal view H lateral view I dorsal view J ventral view. Abbreviations: asa arm spine articulation, d dorsal, de depression, dist distal, lac lateral ambulacral canals, mo muscle opening, no nerve opening, pb podial basin, prox proximal, st secondary teeth, v ventral. Scale bars: 800 Âµm (FâJ); 500 Âµm (C); 300 Âµm (A, B, E); 200 Âµm (D).
	Figure 10. Asteroschemarubrum Lyman, 1879 (IDSSE-EEB-SW0072) A dorsal view B ventral view C dorsal disc D skin on the disc E lateral disc F ventral disc G dorsal arm (proximal) H lateral arm (middle) I ventral arm (proximal) J outer arm spine (proximal) K arm spines (middle) L arm spines (distal). Abbreviations: as adoral shield, ass adoral shield spine, ars arm spine, go granular ossicle, gs genital slit, iars inner arm spine, j jaw, oars outer arm spine, ass adoral shield spine, rs radial shield, t teeth, tp tentacle pore. Scale bars: 2 mm (A, B); 1 mm (EâG, I); 500 Âµm (C, H, J, K); 200 Âµm (D, L).
	Figure 11. Asteroschemarubrum Lyman, 1879 (IDSSE-EEB-SW0072) A, B lateral arm plate (external, internal), C outer arm spine (middle), D arm spine (distal), E skin from dorsal arm base (rounded to somewhat cone shaped granular ossicles), FâJ vertebrae; F proximal view, G distal view, H lateral view, I dorsal view, J ventral view. Abbreviations: d dorsal, de depression, dist distal, lac lateral ambulacral canals, mo muscle opening, no nerve opening, pb podial basin, prox proximal, st secondary teeth, v ventral. Scale bars: 800 Âµm (C); 500 Âµm (A, F, G); 300 Âµm (B, E, HâJ); 200 Âµm (D).
	Figure 12. Asteroschematubiferum Matsumoto, 1911 (IDSSE-EEB-SW0078) A dorsal view B ventral view C dorsal disc D skin on the arm E lateral disc F ventral disc G dorsal arm (proximal) H dorsal arm (middle) I ventral arm (proximal) J outer arm spine (proximal) K arm spines (middle) L arm spines (distal). Abbreviations: as adoral shield, ars arm spine, go granular ossicle, gs genital slit, iars inner arm spine, j jaw, oars outer arm spine, rs radial shield, t teeth, tp tentacle pore. Scale bars: 2 mm (A, B); 1 mm (EâG, I); 500 Âµm (C, H, J, K); 200 Âµm (D, L).
	Figure 13. Asteroschematubiferum Matsumoto, 1911 (IDSSE-EEB-SW0078) A lateral arm plate B outer arm spine (middle) C inner arm spine (middle) D arm spine (distal) E skin from dorsal arm base (granular ossicles) FâJ vertebrae F proximal view G distal view H lateral view I dorsal view J ventral view. Abbreviations: d dorsal, dist distal, lac lateral ambulacral canals, mo muscle opening, no nerve opening, pb podial basin, prox proximal, st secondary teeth, v ventral. Scale bars: 800 Âµm (F, G, J): 500 Âµm (B, H, I); 300 Âµm (A, CâE).
	Figure 14. Asteroschemasalix Lyman, 1879 (IDSSE-EEB-SW0082) A dorsal view B ventral view C dorsal disc D skin on the arm E ventral disc F dorsal arm (proximal) G lateral arm (middle) H ventral arm (proximal) I ventral arm (distal) J outer arm spine (proximal) K arm spines (middle) L arm spines (distal). Abbreviations: as adoral shield, ass adoral shield spine, ars arm spine, go granular ossicle, gs genital slit, iars inner arm spine, j jaw, oars outer arm spine, ots oral tentacle scale, rs radial shield, t teeth, tp tentacle pore. Scale bars: 2 mm (A, B); 1 mm (C, EâH); 500 Âµm (J, K); 200 Âµm (D, I, L).
	Figure 15. Asteroschemasalix Lyman, 1879 (IDSSE-EEB-SW0082) A lateral arm plate B outer arm spine (middle) C inner arm spine (middle) D arm spine (distal) E skin from dorsal arm base (granular ossicles) FâJ vertebrae F proximal view G distal view H lateral view I dorsal view J ventral view. Abbreviations: d dorsal, dist distal, lac lateral ambulacral canals, mo muscle opening, no nerve opening, pb podial basin, prox proximal, st secondary teeth, v ventral. Scale bars: 800 Âµm (B, FâJ); 300 Âµm (A, C, E); 100 Âµm (D).
	Figure 16. Asteroschemacf.lissum H. L. Clark, 1939 (IDSSE-EEB-SW0079) A dorsal view B ventral view C dorsal disc D skin on the disc E lateral disc F ventral disc G dorsal arm (middle) H lateral arm (middle) I ventral arm (proximal) J outer arm spine (proximal) K arm spines (middle) L arm spines (distal). Abbreviations: as adoral shield, ass adoral shield spine, ars arm spine, go granular ossicle, gs genital slit, iars inner arm spine, j jaw, oars outer arm spine, ots oral tentacle scale, rs radial shield, t teeth, tp tentacle pore. Scale bars: 2 mm (A, B, G); 1 mm (C, E, F, H, I); 500 Âµm (J, K); 200 Âµm (D, L).
	Figure 17. Asteroschemacf.lissum H. L. Clark, 1939 (IDSSE-EEB-SW0079) A, B lateral arm plate (external, internal) C outer arm spine (middle) D arm spine (distal) E skin from dorsal arm base (granular ossicles) FâJ vertebrae F proximal view G distal view H lateral view I dorsal view J ventral view. Abbreviations: d dorsal, dist distal, lac lateral ambulacral canals, mo muscle opening, no nerve opening, pb podial basin, prox proximal, st secondary teeth, v ventral. Scale bars: 800 Âµm (FâJ); 500 Âµm (A, C); 300 Âµm (B, D); 200 Âµm (E).
	Figure 18. Asterostegusmaini McKnight, 2003 (AâMIDSSE-EEB-SW0076 NâPIDSSE-EEB-SW0075) A dorsal view B center of dorsal disc C dorsal disc (distal end of radial shields) D ventral disc E, F oral frame G lateral disc H dorsal arm (proximal) I dorsal arm (distal) J lateral arm (proximal) K ventral arm (middle) L, M arm spines (distal) N oral frame O periphery of the disc P oral frame (small transparent ossicles). Abbreviations: as adoral shield, ars arm spine, go granular ossicle, gs genital slit, j jaw, mp median plate, oip oral interradial plate, rs radial shield, t teeth, tp tentacle pore, ts tentacle scale. Scale bars: 6 mm (A); 2 mm (BâH, N); 1 mm (IâK, O, P); 500 Âµm (L); 200 Âµm (M).
	Figure 19. Asterostegusmaini McKnight, 2003 (IDSSE-EEB-SW0076) A, B lateral arm plate (external, internal) C, D arm spine (middle) E, F vertebrae (middle) E proximal view F distal view GâK vertebrae (distal) G proximal view H distal view I lateral view J dorsal view K ventral view. Abbreviations: asa arm spine articular structure, d dorsal, de depression, dist distal, mo muscle opening, ob oral bridge, prox proximal, v ventral. Scale bars: 800 Âµm (E, F); 500 Âµm (A, C, D, I, K); 300 Âµm (B, G, H, J).
	Figure 20. Astrodendrumcf.sagaminum (DÃ¶derlein, 1902) (IDSSE-EEB-SW0104) A dorsal view B ventral view C dorsal disc (center) D dorsal disc (distal edge) E radial shield F, G ventral disc H oral frame I, J ventral view of arm base K ventral view of after second arm branch L, M dorsal view of second arm branch N lateral view of arm (middle) O lateral view of arm (pedicellariae with baseplate) P, Q variations of arm spine on distal end of the arm. Abbreviations: ars arm spine, co conical ossicles gs genital slit, os oral shield, peb = pedicellarial band, rs radial shield, t teeth, tp tentacle pore. Scale bars: 16 mm (A, B); 2 mm (C, D, F, L); 1 mm (E, HâK, M, N); 500 Âµm (G, O); 200 Âµm (P, Q).
	Figure 21. Astrodendrumcf.sagaminum (DÃ¶derlein, 1902) (IDSSE-EEB-SW0104) A lateral arm plate (middle) B plan view of baseplate C pedicellariae D arm spine on distal end EâJ vertebrae on middle half of arm E proximal view of branch vertebrae F proximal view G distal view H lateral view I dorsal view J ventral view KâN vertebrae on distal end of arm K distal view L proximal view M lateral view N ventral view. Abbreviations: ap articular pad of the base, au auricle, d dorsal, dist distal, fo foramina of the base, fs fossa between adjacent tubercles, hd head of the apophysis, lac lateral ambulacral canals, mo muscle opening, no nerve opening, pd pedicel of the apophysis, prox proximal, pt primary tooth of the blade, sh sheath of the baseplate, su sulcus of tubercle head, v ventral. Scale bars: 800 Âµm (EâJ); 500 Âµm (A, B); 300 Âµm (KâN); 200 Âµm (D); 80 Âµm (C).

References [+] :

Boissin, DNA barcoding of reef brittle stars (Ophiuroidea, Echinodermata) from the southwestern Indian Ocean evolutionary hot spot of biodiversity. 2017, Pubmed, Echinobase

﻿Order Euryalida (Echinodermata, Ophiuroidea), new species and new records from the South China Sea and the Northwest Pacific seamounts.

Order Euryalida (Echinodermata, Ophiuroidea), new species and new records from the South China Sea and the Northwest Pacific seamounts.