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Figure 4. Enigmaticolus nipponensis comb. nov. A in Integrative taxonomy of enigmatic deep-sea true whelks in the sister-genera Enigmaticolus and Thermosipho (Gastropoda: Buccinidae)

Figure 4. Enigmaticolus nipponensis comb. nov. A, Natsu site, Iheya North hydrothermal field (27°46.8522'N, 126°54.0584'E), 1078 m deep, shell length 90.4 mm (NSMT-Mo 79173). B, Aki site, Iheya North hydrothermal field (27°46.103'N, 126°54.090'E), 1086 m deep, shell length 84.6 mm (NSMT-Mo 79174). C, Aki site, Iheya North hydrothermal field (27°46.103'N, 126°54.090'E), 1086 m deep, shell length 79.0 mm (NSMT Mol-79175). D, Kuroshima Knoll hydrocarbon seep site (24°07.830'N, 124°11.562'E), 641 m deep, shell length 88.7 mm (NSMT Mol-79176). Scale bars = 2 cm.

Figure 5. A–C, Enigmaticolus nipponensis comb. nov. A in Integrative taxonomy of enigmatic deep-sea true whelks in the sister-genera Enigmaticolus and Thermosipho (Gastropoda: Buccinidae)

Figure 5. A–C, Enigmaticolus nipponensis comb. nov. A, 'F site' methane seep, South China Sea (22°06.945'N, 119°17.128'E), 1128 m deep, shell length 94.9 mm (HKBU Mol-2020010001). B, 'F site' methane seep, South China Sea (22°06.945'N, 119°17.128'E), 1128 m deep, shell length 87.5 mm (HKBU Mol-2020010002). C, 'F site' methane seep, South China Sea (22°6.912'N, 119°17.104'E), 1127 m deep, shell length 91.4 mm (HKBU Mol-2020010003). D, Thermosipho auzendei, holotype (MNHN IM-2000–7049), Rehu site, East Pacific Rise at 17°S (17°24.85'S, 113°12.15'W), 2578 m deep, shell length 62.0 mm (photo courtesy of Philippe Maestrati). Scale bars = 2 cm.

Figure 9. A, B, Enigmaticolus nipponensis comb. nov. A in Integrative taxonomy of enigmatic deep-sea true whelks in the sister-genera Enigmaticolus and Thermosipho (Gastropoda: Buccinidae)

Figure 9. A, B, Enigmaticolus nipponensis comb. nov. A, holotype of Enigmaticolus monnieri (MNHN IM-2000–20464), Tulear, Madagascar, trawled by commercial fishery vessels (shrimpers), Mozambique Channel, 800 m deep, shell length 63.7 mm (photo courtesy of Manuel Caballer). B, MIRIKY station CP3279 (15°22'S, 45°57'E), between Majunga and CapSaint-André, Madagascar, 780–1020 m deep, shell length 99.4 mm (MNHN IM-2009–7079; photo courtesy of Delphine Brabant and Barbara Buge). C, Enigmaticolus marshalli. Holotype (NMNZ M.274915), NIWA 32231 station 2007204 (25°48'S, 177°10'W), Monowai Caldera, south-south-west of Tonga, Kermadec Ridge, New Zealand, 1026 m deep, shell length 90.1 mm. D, Enigmaticolus voluptarius, holotype (MNHN IM-2000–27067), EBISCO station CP2554 (21°05'S, 158°33'E), Bellona, New Caledonia, 704–720 m deep, shell length 77.8 mm. Scale bars = 2 cm.

Figure 7 in Integrative taxonomy of enigmatic deep-sea true whelks in the sister-genera Enigmaticolus and Thermosipho (Gastropoda: Buccinidae)

Figure 7. Radular variation in Enigmaticolus nipponensis comb. nov. A, Natsu site, Iheya North hydrothermal field (27°46.8522'N, 126°54.0584'E), 1078 m deep (NSMT-Mo 79172). B, 'F site' seep, South China Sea (22°06.945'N, 119°17.128'E), 1128 m deep (HKBU Mol-2020010001). Scale bars = 200 μm.

Figure 8 in Integrative taxonomy of enigmatic deep-sea true whelks in the sister-genera Enigmaticolus and Thermosipho (Gastropoda: Buccinidae)

Figure 8. Maximum likelihood molecular phylogenetic tree of Buccinidae based on a 645-bp alignment of the COI gene. Node values are bootstrap values, only those above 70 are shown. Scale bars indicate the number of substitutions per site. Numbers appearing at the end of the names are the GenBank accession numbers. SCS, South China Sea.

Figure 5. A–C, Enigmaticolus nipponensis comb. nov. A in Integrative taxonomy of enigmatic deep-sea true whelks in the sister-genera Enigmaticolus and Thermosipho (Gastropoda: Buccinidae)

Figure 3. Enigmaticolus nipponensis comb. nov. A in Integrative taxonomy of enigmatic deep-sea true whelks in the sister-genera Enigmaticolus and Thermosipho (Gastropoda: Buccinidae)

Figure 3. Enigmaticolus nipponensis comb. nov. A, holotype (NSMT-Mo 71689), Iheya North Original site, Iheya North hydrothermal field (27°47.180'N, 126°54.149'E), 1049 m deep, shell length 93.8 mm (photo courtesy of Kazunori Hasegawa). B, paratype #4 (JAMSTEC 009556), Sumisu Caldera, Izu-Ogasawara Arc (31°47.180'N, 126°54.149'E), 1049 m deep, shell length 90.1 mm (photo courtesy of Takashi Okutani). C, Iheya North Original site, Iheya North hydrothermal field (27°47.4495'N, 126°53.8149'E), 1002 m deep, shell length 80.2 mm (NSMT-Mo 79171). D, Natsu site, Iheya North hydrothermal field (27°46.8522'N, 126°54.0584'E), 1078 m deep, shell length 81.2 mm (NSMT-Mo 79171). Scale bars = 2 cm.

Figure 2 in Integrative taxonomy of enigmatic deep-sea true whelks in the sister-genera Enigmaticolus and Thermosipho (Gastropoda: Buccinidae)

Figure 2. In situ observations of deep-sea whelks as indicated by white arrows. A, Natsu site, Iheya North hydrothermal field (27°46.8522'N, 126°54.0584'E), 1078 m deep, by JAMSTEC ROV Hyper-Dolphin. B, Iheya North Original site, Iheya North hydrothermal field (27°47.4495'N, 126°53.8149'E), 1002 m deep. C, Aki site, Iheya North hydrothermal field (27°46.103'N, 126°54.090'E), 1086 m deep. D, E, in mussel (Bathymodiolus spp.) bed, 'F site', South China Sea (22°06.945'N, 119°17.128'E), 1128 m deep, by ROV ROPOS. F, on a authigenic carbonate cliff near edge of the seep, 'F site', South China Sea (22°6.912'N, 119°17.104'E), by ROV ROPOS.

Figure 1 in Integrative taxonomy of enigmatic deep-sea true whelks in the sister-genera Enigmaticolus and Thermosipho (Gastropoda: Buccinidae)

Figure 1. Map of the north-western Pacific showing the relevant vent and seep sites where new materials were collected for the present study. A, overview. B, close-up of the Izu-Bonin Arc area. C, close-up of the Okinawa Trough area. Triangles indicate hydrothermal vent fields and circles indicate hydrocarbon seep sites. 1, Myojin Knoll vent field; 2, Sumisu Caldera vent site; 3, Iheya North vent field; 4, Kuroshima Knoll seep site; 5, 'F site' seep, South China Sea. Maps generated with Mercator projection using the General Bathymetric Chart of the Oceans (GEBCO; https://www.gebco.net/; date accessed August 13, 2020) grid display software and GEBCO one-arc minute interval grid.

FIGURE 22. A, B. Sulcosinus carinatus n in Seven new genera and thirty-four new species of buccinoid gastropods (Neogastropoda: Buccinidae) from the Aleutian Islands, Alaska

FIGURE 22. A, B. Sulcosinus carinatus n. sp. A. Holotype, LACM 3594, S of Kiska Island, 470 m, 46.8 mm; B. Radula, Holotype, bar = 200 µm; C–E. Sulcosinus taphrium (Dall, 1891); C. Holotype, USNM 122548, N of Akutan Island, 642 m, 42 mm; D. RNC 4000, SW of Saint George Island, 485 m, 45.5 mm; E. Radula, RNC 4000, bar = 200 µm; F, G.–Buccinum lanatum n. sp.; F. Holotype, SBMNH 169021, SW of Herbert Island, 238 m, 42 mm; G. Radula, Holotype, bar = 200 µm; H. Buccinum tunicatum Golikov & Gulbin, 1977, Holotype ZIN 26225/1, Urup Island, Kurile Is., Russia, 48 m, 27 mm; I. Buccinum fringillum Dall, 1877, RNC 4824, Chukchi Sea, 22 m, 23 mm; J, K. Buccinum katharinae n. sp.; J. Holotype, LACM 3595, Amutka Pass, 325 m, 29.2 mm; K. Radula, Holotype, bar = 200 µm.

FIGURE 19. A–C. Volutopsius nanus n in Seven new genera and thirty-four new species of buccinoid gastropods (Neogastropoda: Buccinidae) from the Aleutian Islands, Alaska

FIGURE 19. A–C. Volutopsius nanus n. sp.; A. Holotype, 3588, SW of Buldir Island, 325 m, 35.6 mm; B. Holotype, close-up of sculpture; C. Radula, Paratype, RNC 4711, bar = 200 µm; D, E. Volutopsius gracilis n. sp. D. Holotype, LACM 3589, N of Unimak Island, 80 m, 142.1 mm; E. Radula, Paratype, RNC 5031, bar = 200 µm; F, G. Volutopsius castaneum (Mörch, 1857); F. RNC 4801, Adak Island, 13 m, 76.2 mm; G. Radula, RNC 4801, bar = 200 µm; H, I. Volutopsius stefanssoni Dall, 1919; H. RNC 5038, Chukchi Sea, 42 m, 79.0 mm; I. Radula, RNC 4272, bar = 200 µm; J. Volutopsius stefanssoni var. simplex, RNC 4005, N of Shemya Island, 115 m, 99.6 mm; K–M. Crebrivolutopsius labidentatus n. sp. K. Holotype, CASIZ 137734, Petrel Bank, 213 m, 107.6 mm; L. RNC 4658, close-up of labial teeth, S of Amchitka Island, 388 m, 72.3 mm; M. Radula, RNC 4658, bar = 200 µm.

FIGURE 20. A–E in Seven new genera and thirty-four new species of buccinoid gastropods (Neogastropoda: Buccinidae) from the Aleutian Islands, Alaska

FIGURE 20. A–E. Aleutibuccinum clarki (Kantor & Harasewych, 1998); A. Paratype, RNC 3924, S of Attu Island, 169 m, 50.7 mm; B. RNC 3925, S of Attu Island, 169 m, 50.1 mm; C. RNC 4371, E of Segula Island, 336 m, 55.6 mm; D. RNC 3962, Seguam Pass, 235 m, 61.1 mm; E. operculum from RNC 3924 (fig. 20 A); F–H. Castaneobuccinum castaneum Dall, 1877; F. Holotype, USNM 108973, Shumagin Islands, 36 m, 65 mm; G. RNC 4722, Tanaga Island, 90 m, 69 mm; H. Radula, RNC 4722, bar = 200 µm; I, J. Castaneobuccinum fluctuatum Dall, 1919; I. Holotype, USNM 205916, Bering Sea, off Saint George Island, 54 m, 90 mm; J. RNC 4731, Petrel Bank, 91 m, 78 mm; K, L. Castaneobuccinum triplostephanum Dall, 1919; K. USNM 108974, Kiska Island, 9 m, 75 mm; L. RNC 5054, SE of Buldir Island, 51 m, 63.8 mm.

FIGURE 15. A–C. Neptunea aleutica n in Seven new genera and thirty-four new species of buccinoid gastropods (Neogastropoda: Buccinidae) from the Aleutian Islands, Alaska

FIGURE 15. A–C. Neptunea aleutica n. sp.; A. Holotype, LACM 3580, S of Oglodak, 273 m, 108.5 mm; B. Paratype, RNC 4715, E of Kiska Island, 252 m, 107.3 mm; C. Radula, Paratype, RNC 4715, bar = 200 µm; D–F. Neptunea baxteri n. sp.; D.–Holotype, LACM 3581, N of Umnak Island, 270–296 m, 114 mm; E. Paratype, RNC 4766, NE of Amlia Island, 92 m, 89.5 mm; F. Radula, Paratype, CASIZ 236196, bar = 200 µm. G–J. Neptunea dominator n. sp. G. Holotype, LACM 3582, Petrel Bank, 256 m, 111.6 mm; H. RNC 4673, SW of Tanaga Island, 210 m, 89.0 mm; I. Paratype, RNC 4621, SW of Buldir Island, 325 m, 51.3 mm; J. Radula, holotype, bar = 200 µm.

FIGURE 16. A–D. Neptunea petrelensis n in Seven new genera and thirty-four new species of buccinoid gastropods (Neogastropoda: Buccinidae) from the Aleutian Islands, Alaska

FIGURE 16. A–D. Neptunea petrelensis n. sp.; A. Holotype LACM 3583, Petrel Bank, 143 m, 74.5 mm; B. Paratype, RNC 4984, Petrel Bank, 101 m, 102.8 mm; C. Radula, paratype, RNC 4984, bar = 200 µm; D–F. Neptunea quhmax n. sp.; D. Holotype, LACM 3584, Stalemate Bank, 219 m, 109.5 mm; E. Paratype, SBMNH 169009, Stalemate Bank, 222 m, 97.4 mm; F. Radula, Paratype, RNC 4599, Stalemate Bank, bar = 200 µm; G, H. Neptunea alabaster Alexeyev & Fraussen, 2005; G. Paratype, RNC 4069, Bering Sea, W of Cape Olyutor, Russia, 290 m, 113.8 mm; H. Radula, RNC 4464, Bering Sea, Pervenets Canyon, 913 m, bar = 200 µm; I–K. Neptunea vesteraalen n. sp.; I. Holotype, LACM 3585, W of Akutan Island, 411 m, 160 mm; J. Paratype 3, USNM 1606669, S of Herbert Island, 238 m, 127 mm; K. Radula, paratype, RNC 4510, bar = 200 µm.

FIGURE 13. Exiloberingius fragilis n in Seven new genera and thirty-four new species of buccinoid gastropods (Neogastropoda: Buccinidae) from the Aleutian Islands, Alaska

FIGURE 13. Exiloberingius fragilis n. sp., A. Holotype, LACM 3579, SW of Seguam Island, 153 m, 106.6 mm; B. Paratype, RNC 4877 (type locality), 78.5 mm; C, Paratype, LACM 3611, S of Seguam Island, 411 m, 109.5 mm (fragmented); D. Radula, paratype 1, bar = 200 µm.

FIGURE 14. A–D. Aulacofusus canaliculatus n in Seven new genera and thirty-four new species of buccinoid gastropods (Neogastropoda: Buccinidae) from the Aleutian Islands, Alaska

FIGURE 14. A–D. Aulacofusus canaliculatus n. sp. A. Holotype, LACM 3667, SE of Agattu Island, 135 m, 66.6 mm; B. Paratype, RNC 4143, S of Buldir Island, 71 m, 53.3 mm; C. Aulacofusus periscelidus (Dall, 1891), RNC 4860, W of Hawadax (Rat) Island, 94 m, 77.47 mm; D. RNC 4625, Adak Island, 18 m, 69.1 mm; E–G. Aulacofusus tanagaensis n. sp.; E. Holotype, LACM 3665, W of Tanaga Island, 53 m, 54.9 mm; F. Paratype, RNC 4585, W of Tanaga Island, 95 m, 54.0 mm; G. Radula, paratype, bar = 200 µm.

FIGURE 6. A in Seven new genera and thirty-four new species of buccinoid gastropods (Neogastropoda: Buccinidae) from the Aleutian Islands, Alaska

FIGURE 6. A. Metajapelion adelphicus (Dall, 1907) RNC 4967, Off Okinoshima Island, Japan, 182 m, 97.4 mm; B–E. Aleutijapelion aleuticus (Dall, 1895); B. Holotype, USNM 106999, Amutka Pass, 454 m, 65.0 mm; C. RNC 4772, S of Umnak Island, 86 m, 74.3 mm; D. Operculum of RNC 4772, 28 mm; E. Radula, RNC 4880, W of Carlisle Island, 228 m, bar = 100 µm; F–H. Aleutijapelion mirandus; F. Holotype, LACM 3664, N of Amlia Island, 423 m, 74.4 mm; G. RNC 5019, E of Kiska Island, 268 m, 91.7 mm; H. Radula, RNC 4639, S of Atka Island, 207 m, bar = 200 µm.

FIGURE 7. A–D. Beringius nearensis n in Seven new genera and thirty-four new species of buccinoid gastropods (Neogastropoda: Buccinidae) from the Aleutian Islands, Alaska

FIGURE 7. A–D. Beringius nearensis n. sp.; A. Holotype, LACM 3571, Stalemate Bank, W of Attu, 219 m, 103.4 mm; B. Paratype 4, RNC 4882, E of Agattu Island, 111 m, 85.8 mm; C. Paratype, USNM 1606662, 127.1 mm; D. Radula, Paratype, CASIZ 236197; E–F. Beringius undatus Dall, 1919; E. RNC 4745, N of Akutan Island, 92 m, 129.6 m; F. Radula, RNC 4745, bar = 200 µm; G–H. Beringius crebricostatus (Dall, 1877); G. RNC 4009, Unalaska Island, 5 m, 90.5 mm; H. Radula, RNC 4009, bar = 200 µm.

FIGURE 9. A–C. Beringius amliensis n in Seven new genera and thirty-four new species of buccinoid gastropods (Neogastropoda: Buccinidae) from the Aleutian Islands, Alaska

FIGURE 9. A–C. Beringius amliensis n. sp.; A. Holotype, LACM 3572, N of Amlia Island, 149 m, 120.7 mm; B. Paratype 4, RNC 4502, N of Amlia Island, 72 m, 116.2 mm; C. Radula, Holotype, bar = 200 µm; D–G. Beringius incisus (Dall, 1907); D. Holotype, USNM 110488, Petrel Bank, 95 m, 75 mm; E. RNC 4463, Atka Island, 73 m, 93.8 mm; F. RNC 4883, Petrel Bank, 81 m, 72.4 mm; G. Radula, RNC 4883, bar = 200 µm.

FIGURE 10. A–D. Beringius bisulcatus n in Seven new genera and thirty-four new species of buccinoid gastropods (Neogastropoda: Buccinidae) from the Aleutian Islands, Alaska

FIGURE 10. A–D. Beringius bisulcatus n. sp.; A. Holotype, LACM 3573, Tanaga Island, 95 m, 100.9 mm; B. Paratype 3, RNC 4475, S of Gareloi Island, 182 m, 115.0 mm; C. RNC 4802, Tanaga Island, 95 m, 42.0 mm; D. Radula, RNC 4801, bar = 200 µm; E–G. Beringius kiskensis n. sp. E. Holotype, LACM 3574, Kiska Island, 97 m, 85.4 mm; F. Paratype, RNC 4910, E of Kiska Island, 96 m, 112.7 mm; G. Radula, RNC 4910, bar = 200 µm.