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FIGURE 16. Frontohornera frontalis n. gen., n in Bryozoa (Cyclostomata and Ctenostomata) from polymetallic nodules in the Russian exploration area, Clarion - Clipperton Fracture Zone, eastern Pacific Ocean-taxon novelty and implications of mining

FIGURE 16. Frontohornera frontalis n. gen., n. sp. Holotype specimen, ZIRAS 1/50707, respectively seen in A, frontal and B, abfrontal views. Scale bars: 1 mm.

FIGURE 15. Abyssoecia elevata n. gen., n in Bryozoa (Cyclostomata and Ctenostomata) from polymetallic nodules in the Russian exploration area, Clarion - Clipperton Fracture Zone, eastern Pacific Ocean-taxon novelty and implications of mining

FIGURE 15. Abyssoecia elevata n. gen., n. sp. Progressive stages of development of young ancestrulate colonies viewed from different angles, showing development of either supportive columns or growth of lateral margins towards substratum as colony grows larger. A–C, specimen GLD4–09, Stn 196; D–F, specimen YMG4–07, Stn 129; G–L, specimen YMG4–04, Stn 44. Scale bars: A, B, D–F, 250 µm; C, 200 µm; G–L, 500 µm.

FIGURE 13. Abyssoecia elevata n. gen., n in Bryozoa (Cyclostomata and Ctenostomata) from polymetallic nodules in the Russian exploration area, Clarion - Clipperton Fracture Zone, eastern Pacific Ocean-taxon novelty and implications of mining

FIGURE 13. Abyssoecia elevata n. gen., n. sp. Holotype and paratype colonies detached from their substrata. A, B, holotype, ZIRAS 1/50703, frontal and profile views; note foramen in colony formed by growth and fusion of lobes around a space; C, D, paratype 1, ZIRAS 2/50704 frontal and abfrontal views; E, F, paratype 2, ZIRAS 3/50705, frontal and profile views. Scale bars: 500 µm.

FIGURE 9 in Bryozoa (Cyclostomata and Ctenostomata) from polymetallic nodules in the Russian exploration area, Clarion - Clipperton Fracture Zone, eastern Pacific Ocean-taxon novelty and implications of mining

FIGURE 9. Tubuliporina sp. incertae sedis. Specimen ZIRAS 1/50674. A–C, rotational and D, tilted, views of sole colony; E–H, rotational views of distal part of colony showing two autozooidal peristomes and a smaller-diameter (?female) peristome; I, J, close-ups of smaller peristome; K, close-up of right-hand peristomial aperture in D. Scale bars: A–D, 250 µm; E–H, 200 µm; I–K, 100 µm.

FIGURE 8. Orectopora flabellum n. gen., n in Bryozoa (Cyclostomata and Ctenostomata) from polymetallic nodules in the Russian exploration area, Clarion - Clipperton Fracture Zone, eastern Pacific Ocean-taxon novelty and implications of mining

FIGURE 8. Orectopora flabellum n. gen., n. sp. Colonies at early stages of astogeny. A–D, rotational views of specimen GLD4–08, Stn 151, with ancestrular region more magnified in D; E, F, rotational views of specimen GLD4–08, Stn 145; G, H, specimen GLD4–11, Stn 230; I, J, specimen GLD4–11, Stn 219; K–P, rotational views of paratype specimen, NIWA 127723. Scale bars: A–C, E–H, 300 µm; D, P, 200 µm; I–O, 500 µm.

FIGURE 1 in Bryozoa (Cyclostomata and Ctenostomata) from polymetallic nodules in the Russian exploration area, Clarion - Clipperton Fracture Zone, eastern Pacific Ocean-taxon novelty and implications of mining

FIGURE 1. Sampling stations in the eastern Russian exploration area of the CCZ, ranging from depths of 4510 m (orange) to 5280 m (purplish-blue).

FIGURE 4. Pandanipora helix n. gen., n in Bryozoa (Cyclostomata and Ctenostomata) from polymetallic nodules in the Russian exploration area, Clarion - Clipperton Fracture Zone, eastern Pacific Ocean-taxon novelty and implications of mining

FIGURE 4. Pandanipora helix n. gen., n. sp. Details of morphology. A, specimen YMG4–13, Stn 289; B–D, paratype 4, ZIRAS 5/50671; E, paratype 5, ZIRAS 6/50672; F, G, Q, paratype 1, ZIRAS 2/50668; H, specimen GLD4–11, Stn 217; I, holotype, ZIRAS 1/50667; J, specimen YMG4–06, Stn 68; K, O, specimen YMG4–13, Stn 321; L, specimen YMG4–13, Stn 295; M, paratype 2, ZIRAS 3/50669; N, specimen GLD4–12, Stn 258; P, specimen GLD4–12, Stn 255. A, distal view of growing edge of colony showing early differentiation of daughter zooid and prop from the floor of the parent zooid, with chambers arranged in stacks; B, growing edge of colony showing differentiation of a daughter zooid (lower) from the floor of the parent zooid (upper); C, same, close-up; D, interior surface of developing zooid chamber showing imbricated foliated fabric of wedge-shaped crystallites and openings of pseudopores; E–I, variability of zooid size and geometry: E, elongated zooids with tall peristomes; F, part of ring-like colony; G, lateral view of same colony, two zooids near-perpendicular to plane of props; H, arcuate colony with overlapping peristome bases; I, lateral view of apical coil of spiral colony with subconnate peristome bases and shortened peristomes, most broken; J, frontolateral view of tubular peristome showing annular striae on the surface; K, lateral view of tubular peristome; L, peristomial aperture, frontal view; M, oblique view of peristomial aperture showing imbricated foliated fabric of wedge-shaped crystallites and interior openings of pseudopores; N, imbricated foliated fabric of wedge-shaped crystallites around internal opening of pseudopore; O, zooidal wall showing tiny sparse simple pseudopores; P, Q, close-up of zooidal wall showing crystallites. Scale bars: A, B, G, 250 µm; C, J, K, O, 100 µm; D, N, 10 µm; E, F, H, I, 500 µm; L, M, Q, 50 µm; P, 15 µm.

FIGURE 14. Abyssoecia elevata n. gen., n in Bryozoa (Cyclostomata and Ctenostomata) from polymetallic nodules in the Russian exploration area, Clarion - Clipperton Fracture Zone, eastern Pacific Ocean-taxon novelty and implications of mining

FIGURE 14. Abyssoecia elevata n. gen., n. sp. A–E, paratype 3, ZIRAS 4/50706; F, specimen YMG4–06, Stn 106; G, J, specimen YMG4–13, Stn 302; H, I, N, specimen YMG4–07, Stn 129; K, specimen GLD4–09, Stn 191; L, M, specimen YMG4–04, Stn 44. A, gonozooid at upper left of image with projecting ooeciostome; B, D, distal and distolateral views of gonozooid; C, E, frontal and distal views of ooeciostome with subcircular ooeciopore; F–H, distolateral (F) and distal views of colony margin with developing zooids; I–K, frontal views of short autozooidal peristomes; L–N, ancestrular region of two colonies. Scale bars: A, 500 µm; B, D, F–J, L–N, 200 µm; C, E, 100 µm; K, 50 µm.

FIGURE 12. Discantenna metallica n in Bryozoa (Cyclostomata and Ctenostomata) from polymetallic nodules in the Russian exploration area, Clarion - Clipperton Fracture Zone, eastern Pacific Ocean-taxon novelty and implications of mining

FIGURE 12. Discantenna metallica n. sp. Progressive stages of development of ancestrulate and young colonies. A–D, specimen YMG4–14, Stn 358; E–H, specimen GLD4–11, Stn 224; I–L, specimen GLD4–11, Stn 207. A–D, rotational views (respectively frontal, distolateral, lateral, proximal) of newly formed colony with ancestrular zooid and developing daughter peristome; note developing kenozooids in B; E–H, similar views of a slightly older colony with further kenozooidal development in F; I–L, apical and profile views of an older colony (ancestrular protoecium missing) with a definite stalk underway and peristomes opening all around it. Scale bars: A–H, J, 100 µm; I, K, L, 200 µm.

FIGURE 11. Discantenna metallica n in Bryozoa (Cyclostomata and Ctenostomata) from polymetallic nodules in the Russian exploration area, Clarion - Clipperton Fracture Zone, eastern Pacific Ocean-taxon novelty and implications of mining

FIGURE 11. Discantenna metallica n. sp. Paratype and other colonies. A, B, D, F, G, paratype 2, ZIRAS 3/50702; C, E, H, I, paratype 1, ZIRAS 2/50676; J, L, N, P, specimen YMG4–06, Stn 69; K, M, O, Q, specimen GLD4–08, Stn 151. A, fertile colony with broken base and small capitulum; B, D, same, showing gonozooid in frontal and apertural views; C, E, a second gonozooid with longer ooeciostome; F, G, high- and lower-magnification views of terminal diaphragm, showing skeletal microstructure; H, I, ooeciopores of gonozooid in C and E; J, N, lateral and apical views of colony with small lobate capitulum; K, O, lateral and apical views of another colony with a small capitulum that is more symmetrical; L, M, parts of stalks with short autozooidal peristomes and terminal diaphragms; P, Q, proximal parts of two colonies in post-ancestrular region. Scale bars: A, J, K, N, O, 250 µm; B–E, L, M, P, Q, 100 µm; F–I, 25 µm.

FIGURE 10. Discantenna metallica n in Bryozoa (Cyclostomata and Ctenostomata) from polymetallic nodules in the Russian exploration area, Clarion - Clipperton Fracture Zone, eastern Pacific Ocean-taxon novelty and implications of mining

FIGURE 10. Discantenna metallica n. sp. Holotype and paratype colonies. A–D, holotype, ZIRAS 1/50675; E–I, paratype 1, ZIRAS 2/50676. A, apical and B, lateral views of colony; C, D, same, edge of disk, respectively showing peristomes and differentiating zooids; E, lateral view of colony showing numerous stalk zooids; F, autozooidal peristome near top of stalk with calcified terminal diaphragm; G, underside of part of capitulum seen in E; H, part of stalk with short peristomes and terminal diaphragms; I, margin of capitulum seen in E. Scale bars: A–E, 500 µm; F, 150 µm; G–I, 250 µm.

FIGURE 5. Pandanipora helix n. gen., n in Bryozoa (Cyclostomata and Ctenostomata) from polymetallic nodules in the Russian exploration area, Clarion - Clipperton Fracture Zone, eastern Pacific Ocean-taxon novelty and implications of mining

FIGURE 5. Pandanipora helix n. gen., n. sp. A, K, specimen YMG4–13, Stn 289; B, C, specimen YMG4–13, Stn 308; D, specimen YMG4–07, Stn 143; E, specimen GLD4–11, Stn 219; F, specimen YMG4–14, Stn 330; G–I, specimen YMG4–14, Stn 326; J, specimen GLD4–12, Stn 258; L, specimen YMG4–13, Stn 321. A–D, F, prop morphology: A, vertical and oblique columnar props attached to nodule by expanded tips; B, curved props attached to nodule irregularities by expanded tips with complex outlines; C, same, close-up; D, single elongated prop from its inception in the zooid basal wall to its distal expansion; E, light micrograph of colony showing successively overlapping zooids with their coelomic cavities continuing into the props with no intervening septum or pore, as well as 2–5 brown bodies located centrally to proximally in several zooids; F, prop tip showing opening towards substratum and internal surface with wedge-shaped crystallites, structurally analogous to peristome; G–I, ancestrula and daughter zooid budded from peristome; J–L, ancestrula and postancestrular zooids. Scale bars: A, B, E, 500 µm; C, D, J–L, 250 µm; F, 50 µm; G–I, 100 µm.

FIGURE 7. Orectopora flabellum n. gen., n in Bryozoa (Cyclostomata and Ctenostomata) from polymetallic nodules in the Russian exploration area, Clarion - Clipperton Fracture Zone, eastern Pacific Ocean-taxon novelty and implications of mining

FIGURE 7. Orectopora flabellum n. gen., n. sp. Details of colony morphology. A, B, E, H, holotype colony; C, D, F, G, I, J, specimen GLD4–08, Stn 151. A, B, E, apical, oblique-abfrontal and abfrontal views, respectively, of middle lobe (fascicle) in Fig. 7E, showing apparent roof forming in axil between what will become two fascicle lobes; note relatively large pseudopores in neanic calcification of abfrontal wall in B and E; C, F, two views of a small fascicle showing autozooidal apertures adjacent to concave frontal face of lobe (on right), between which are larger developing chambers (either autozooidal or reproductive), and, along convex abfrontal face (on left), a row of mostly kenozoidal openings (and perhaps some incipient autozooid buds); D, G, J, magnified sequence of images showing the largest developing chamber in C and F, and ultrastructural wall fabric; H, reversed (frontal) view of fascicle tip in E; I, frontal view of chamber in D, G, J. Scale bars: A, B, E, H, 200 µm; C, F, 150 µm; D, I, 50 µm; G, J, 20 µm.

FIGURE 3. Pandanipora helix n. gen., n in Bryozoa (Cyclostomata and Ctenostomata) from polymetallic nodules in the Russian exploration area, Clarion - Clipperton Fracture Zone, eastern Pacific Ocean-taxon novelty and implications of mining

FIGURE 3. Pandanipora helix n. gen., n. sp. Holotype and paratype colonies. A, C, holotype, ZIRAS 1/50667; B, paratype 1, ZIRAS 2/50668; D, paratype 2, ZIRAS 3/50669; E, paratype 3, ZIRAS 4/50670; F, paratype 4, ZIRAS 5/50671; G, paratype 5, ZIRAS 6/50672. A, apical, and C, lateral views of the spiral holotype colony with widened apical part of axis of zooids with mostly broken, shortened peristomes and supported by rare, distanced, elongated tubular props; B, ring-like colony showing attenuated zooids with elongated, gently arcuate tubular peristomes tilted outwards, and supported by strongly elongated, slightly crooked filiform props; note one prop at lower left bifurcated near termination; D, crescentic colony showing zooids with elongated, arcuate tubular peristomes angled outwards, and supported by shortened straight props; E, colony bifurcated near point of origin showing linear chain of zooids with straight peristomes; supported by short props, some with widened terminations; F, arcuate colony with thicker zooids having widened tubular peristomes, supported by gradually shortening props; G, straight colony with greatly elongated peristomes, supported by regular series of props with expanded terminal bases, including four attached to piece of nodule. Scale bars: 500 µm.

FIGURE 2 in Bryozoa (Cyclostomata and Ctenostomata) from polymetallic nodules in the Russian exploration area, Clarion - Clipperton Fracture Zone, eastern Pacific Ocean-taxon novelty and implications of mining

FIGURE 2. Colonies of some cyclostome bryozoans, in vivo, attached to polymetallic nodules. A–E, Pandanipora helix n. gen., n. sp.: A, specimen GLD4–09, Stn 190; B, specimen GLD4–12, Stn 262; C, specimen YMG4–07, Stn 143; D, specimen YMG4–13, Stn 295; E, specimen GLD 4–11, Stn 212. F, Tubuliporina sp. indet., specimen YMG18–01, Stn 7. G, H, Abyssoecia elevata n. gen., n. sp.: G, specimen GLD4–09, Stn 196; H, specimen GLD4–09, Stn 191. I, Discantenna metallica n. sp.: specimen GLD4–11, Stn 224. J, K, Frontohornera frontalis n. gen., n. sp.: J, specimen YMG4–07, Stn 124; K, specimen GLD4–11, Stn 210. L, Alyonushka hystricosa n. gen., n. sp.: specimen GLD4–09, Stn 199. M, Calyssopora volcano n. gen., n. sp.: specimen YMG18–01, Stn 33. N, O, Anyuta anastema n. gen., n. sp.: N, specimen GLD4–09, Stn 180; O, specimen YMG4–06, Stn 71. Scale bars: 1 mm.

FIGURE 6. Orectopora flabellum n. gen., n in Bryozoa (Cyclostomata and Ctenostomata) from polymetallic nodules in the Russian exploration area, Clarion - Clipperton Fracture Zone, eastern Pacific Ocean-taxon novelty and implications of mining

FIGURE 6. Orectopora flabellum n. gen., n. sp. Holotype, ZIRAS 1/50673. A, C, E, colony respectively seen in frontal, abfrontal and apical views; B, frontal view of left-hand lobe (fascicle) in A; D, oblique abfrontal view of right-hand lobe (fascicle) in C; F, base of broken colony stalk. Scale bars: A, C, E, 500 µm; B, D, 250 µm; F, 150 µm.

Figure 7 in Taxonomy of the bryozoan genera Oncousoecia, Microeciella and Eurystrotos (Cyclostomata: Oncousoeciidae)

Figure 7. Microeciella suborbicularis (Hincks, 1880). (A, B, C, D, G) Lectotype (NHM 99.5.1.1427a); (A) overgrown colony origin; (B) autozooids and three gonozooids; (C) typical gonozooid; (D) aberrant gonozooid with subterminal ooeciopore (G) autozooidal frontal wall showing pseudopores. (E, F) gold-coated specimen figured by Hayward and Ryland (1985b, Figure 32A) as Eurystrotos compacta (Norman, 1867); (E) subcircular colony; (F) gonozooid. Scale bars: 1 mm (F); 500 mm (A, B); 200 mm (C, D, E); 50 mm (G).

Figure 4 in Taxonomy of the bryozoan genera Oncousoecia, Microeciella and Eurystrotos (Cyclostomata: Oncousoeciidae)

Figure 4. Oncousoecia dilatans (Johnston, 1847). (A, C, D) Lectotype of Alecto compacta Norman, 1867 (NHM 11.10.1.63); (A) oligoserial branches close to colony origin; (C) gonozooid; (D) another gonozooid illustrating within-colony variability in shape. (B, G) Colony sharing the same substrate as the lectotype (NHM 11.10.1.63); (B) mass of branches overgrowing colony origin; (G) frontal wall of autozooids showing pseudopores and a larger boring. (E, F) Bleached topotype colony (NHM 11.10.1.64); (E) gonozooid; (F) two branches with intact and broken gonozooids. Scale bars: 1 mm (A); 500 mm (B, F); 200 mm (C, D, E); 50 mm (G).

Figure 5 in Taxonomy of the bryozoan genera Oncousoecia, Microeciella and Eurystrotos (Cyclostomata: Oncousoeciidae)

Figure 5. Comparison of early astogeny in (A–C) Alecto compacta (Norman, 1867) [5Oncousoecia dilatans (Johnston, 1847)] (NHM 11.10.1.64) and (D–F) Microeciella suborbicularis (Hincks, 1880), two species formerly considered synonymous. (A) Ancestrula and first budded zooids; (B) ancestrula with short distal tube; (C) large protoecium with tiny pseudopores around perimeter (best seen top right); (D) fan-shaped young colony; (E) ancestrula with long distal tube; (F) protoecium with a few, non-perimetrical pseudopores and two larger, enigmatic punctures or borings (left). Scale bars: 500 mm (D); 200 mm (A); 100 mm (B, E); 50 mm (C, F).

Figure 2. Oncousoecia lobulata Canu, 1918 in Taxonomy of the bryozoan genera Oncousoecia, Microeciella and Eurystrotos (Cyclostomata: Oncousoeciidae)

Figure 2. Oncousoecia lobulata Canu, 1918, lectotype (NHM 1911.10.1.86). (A) Lobate, oligoserial branches; (B) autozooids; (C) gonozooid; (D) gonozooidal pseudopores showing partial occlusion by spines. Scale bars: 1 mm (A); 200 mm (B, C); 50 mm (D).