Bradabyssa mammillata (Grube, 1877) Salazar-Vallejo, 2017

Salazar-Vallejo, Sergio I., 2017, Revision of Brada Stimpson, 1853, and Bradabyssa Hartman, 1967 (Annelida, Flabelligeridae), Zootaxa 4343 (1), pp. 1-98 : 52-54

Bradabyssa mammillata (Grube, 1877) n. comb.

Figure 26

Brada mammillata Grube, 1877a: 69, 1877b: 541 .—McIntosh 1885: 370–371, Pl. 43, Fig. 11 , Pl. 23A, Figs 7 , 8 .—Ehlers 1897: 109.—Fauvel 1932: 185–186, 1953: 352.—Monro 1939: 131.—Hartman 1953: 51.—Hartman 1966: 33 Pl. 9, Figs 3 , 4 .—Hartman 1967: 122.—Wiktor 1980: 277.—Rozbaczylo 1985: 158.— Hartwich 1993: 116.

Type material. Southern Indian Ocean. Kerguelen Islands. One syntype ( MNHW 264), Successful Bay, (Kerguelen Islands in Grube 1877b:511), R.V. Gazelle, Schleinitz or Buchholz, coll. (damaged, most larger papillae eroded, body wall ventrally broken by chaetigers 5–8; 50 mm long, 6.5 mm wide, cephalic cage 4 mm long, 31 chaetigers). Four syntypes ( ZMB 688), complete, Whale Bay, R.V. Gazelle, Schleinitz or Buchholz coll. (10–67 mm long, 4.5–10.0 mm wide, cephalic cage 2–5 mm long, 27–30 chaetigers). Two syntypes ( ZMB 689), off Mount Campbell, R.V. Gazelle, Schleinitz or Buchholz coll. (complete syntype 51 mm long, 8 mm wide, cephalic cage 3 mm long, 29 chaetigers; tubercles large, rounded, arranged in 3 transverse series, anterior ones closer to each other, median series with larger tubercles; larger dorsal tubercles arranged in 4 longitudinal series; ventral papillae much smaller, arranged in 5 transverse series with alternating papillae in successive rows).

Additional material. Southern Indian Ocean. Kerguelen Islands. Three specimens ( MNHN 186 ), complete, 16–18 Jan. 1909 , Rallier du Baty, coll. (50–70 mm long, 7.5–12.0 mm wide, cephalic cage 6–7 mm long, 28–30 chaetigers). Three specimens (MNHN 186a), complete, R.V. Curieuse, C-22, 1914, Rallier du Baty & Soranchet, coll. (one mature female with anterior end exposed; 42.0– 67.5 mm long, 7.5–12.0 mm wide, cephalic cage 4–6 mm long, 30 chaetigers; oocytes 150–200 µm). Five specimens (MNHN-354), only one juvenile complete, with anterior end exposed, Morbihan Bay , 25 Sep. 1951, P . Paulian, coll. (22 mm long, 5 mm wide, cephalic cage 3 mm long, 26 chaetigers). Two specimens ( USNM 4828 ), R.V. Challenger, Sta. 149, Betsy Cove, 46–55 m (most larger tubercles eroded; one with a longitudinal dissection and most viscerae removed; not dissected specimen 67 mm long, 10 mm wide, cephalic cage 1 mm long, 28 chaetigers; gonopodial lobes in chaetiger 5). Antarctica. One specimen ( USNM 56658 ), damaged , USNS Eltanin, Bransfield Strait, Sta. 432 (62°52' S, 59°27' W), 935– 884 m, 7 Jan, 1963 (60 mm long, 6 mm wide, cephalic cage 3 mm long, 30 chaetigers; gonopodial lobes in chaetiger 5). One specimen ( USNM 56664 ), damaged, off South Orkney Island , USNS Eltanin, Sta. 1078 (61°27' S, 41°55' W), 604 m, 12 Apr. 1964. One specimen ( LACM 9550 ), off South Orkney Island , USNS Eltanin, Sta. 1079 (61°26'S, 41°55' W → 61°24' S, 41°55' W), 593–598 m, 13 Apr. 1964. Two specimens ( USNM 56665 ), off South Orkney Island , USNS Eltanin, Sta. 1079 (61°26' S, 41°55' W), 593–598 m, 13 Apr. 1964 (43–101 mm long, 8–11 mm wide, cephalic cage 3 mm long, 31–35 chaetigers; gonopodial lobes in chaetiger 5).

Description. Syntypes (ZMB 689) fusiform, oval in cross section, incurved, slightly to markedly narrower posteriorly ( Fig. 26A ); up to 67 mm long (USNM 56665, 101 mm long), 4.5–10 mm wide, cephalic cage 2–5 mm long (other specimens with chaetae broken), 27–30 chaetigers (USNM 56664, 36 chaetigers). Dorsal tubercles large, globular, in three transverse series per chaetiger, middle series with 5–6 larger, stiff, other series with tubercles less than half as large ( Fig. 26B ); ventral papillae in three rows, rounded (eroded in several syntypes); without tunic, sediment grains adherent to papillae and body tegument.

Cephalic hood not exposed. Anterior end observed by dissection and in non-type specimen (MNHN 186). Prostomium low cone, anterior eyes poorly defined; posterior ones in low rounded lobes. Caruncle extended posteriorly separating branchial plate, median keel pale, lateral ridges dark ( Fig. 26D ). Palps folded; palp keels well developed, rounded.

Branchiae cirriform, sessile on branchial plate, arranged in irregular rows, over 100 per lateral group, some filaments longer than palps. Lateral lips well developed, thick; dorsal lip pigmented, reduced; ventral lip not seen. Nephridial lobes distinct, pale brown, tubular, separated from branchial plate.

Cephalic cage chaetae as long as 1/5–½ body width; chaetae of chaetigers 1–3 slightly longer, directed forward. Chaetae arranged in short lateral series, about 6 capillary chaetae per ramus.

Anterior dorsal margin of first chaetiger with two large rounded papillae. Chaetigers 1–3 of similar length, parapodial development similar. Chaetal transition from cephalic cage to body chaetae abrupt; aristate neurospines present from chaetiger 2. Gonopodial lobes black, in chaetiger 5 ( Fig. 26C , one syntype with a small accessory papilla in chaetiger 6).

Parapodia well developed, lateral. Median neuropodia ventrolateral. Notopodia and neuropodia close to each other. Notopodia less developed than neuropodia, anterior chaetigers with a large trifid rounded infrachaetal papilla, median chaetigers with papillae bifid, posterior chaetigers with single rounded or no infrachaetal papilla. Neuropodia projecting rounded lobes, 3–4 rounded marginal papillae.

Median notochaetae arranged in longitudinal short series; all notochaetae multiarticulate capillaries, most chaetae with short articles basally and medially, medium-sized or long distally, one chaetae with long articles basally ( Fig. 26E ), 4–6 per ramus, 1/6 as long as body width. Aristate neurospines from chaetiger 2, most broken, arranged in J-shaped patterns, 4–5 per ramus ( Fig. 26F ); neurochaetae with very short rings, tips broken, visible in juvenile or smaller specimens ( Fig. 26G ).

Posterior end tapered; pygidium blunt cone, anus terminal, anal cirri absent.

Variation. The different size ranges in the type material shows that the tough body, provided with large globular dorsal tubercles, is a stable feature. The shortest syntype (ZMB-688) is 10 mm long, and shows these features, even though it was fixed in alcohol. Syntypes were 10–67 mm long, 4.5–10.0 mm wide, cephalic cage 2–5 mm long, 27–31 chaetigers. Additional specimens 22–101 mm long, 5–12 mm wide, cephalic cage 3–7 mm long, 26–35 chaetigers.

Remarks. Bradabyssa mammillata (Grube, 1877) n. comb. resembles B. verrucosa (Chamberlin, 1919) n. comb. by having tubercles of different sizes. They differ because B. mammillata has chaetigers 1–3 equally developed whereas in B. verrucosa chaetiger 2 projects more than surrounding chaetigers. One specimen (MNHN 186) has long, pale tubular epibionts; others contain oocytes.

Bradabyssa mammillata also resembles B. elinae by having a single series of larger globular tubercles per segment; however, they differ because B. mammillata has a stiff, wider body, whereas B. elinae is thinner and softer, and because in B. mammillata the anchylose, aristate neurospines have very short rings, whereas in B. elinae they are medium-sized.

Wehe & Fiege (2002:48) stated that there was no type locality indicated for this species; however, the type locality was indicated in the original publication (Grube 1877a:69). The records for the Arabian of Persian Gulf by Wesenberg-Lund (1949:334–335), or from Japan by Imajima (1964:246–247, Figs. 33–38 ), are questionable.

Distribution. Antarctic Ocean and nearby localities, in moderate to abyssal depths.

Figures

FIGURE 3. Brada granosa Stimpson, 1853. A. Juveniles (USNM 453). B. Non-type specimens (USNM 22851), chaetiger 2, right parapodium, anterior view. C. Same, notochaetal basal, medial and distal regions. D. Same, neurochaetal tips. E. Chaetiger 17, right parapodium, anterior view. F. Same, neurochaetal tips. Scale bars: A: 0.8 mm, B, E: 0.3 mm, C: 80 µm, D, F: 20 µm.

FIGURE 4. Brada granulosa Hansen, 1880, syntypes (ZMUB 2197). A. Largest and smallest syntypes, oblique lateral view, anterior end pointing upwards. B. Largest syntype, anterior end, lateral view (arrow points to gonopodial lobe). C. Another syntype, head, frontal view, branchiae and palps removed (Br: branchial scar, PS: palp scar). D. Same, chaetiger 10, right parapodium, posterior view. E. Same, basal, medial and distal notochaetal regions. F. Chaetiger 3, right parapodium, neurochaetae tips (insets: chaetal tips). G. Chaetiger 11, neurochaetae (inset: chaetal tip). Scale bars: A: 6 mm, B: 0.7 mm, C: 0.5 mm, D: 0.1 mm, E: 40 µm, F: 0.2 mm, G: 110 µm.

FIGURE 7. Brada kudenovi n. sp. A. Holotype (CAS 157753), oblique ventral view. B. Same, anterior end (arrow points gonopodial papilla). C. Paratype (CAS 158549), head, frontal view, greenish after methyl-green staining (palps and branchiae removed; Br: branchial scar, PS: palp scar). D. Holotype (CAS 157753), anterior parapodia, chaetigers 4 – 7 (anterior end to the right), oblique ventral view. E. Paratype (CAS 157752), chaetiger 11, right parapodium, anterior view. F. Same, basal, medial and distal notochaetal regions. G. Same, neurochaetae (inset: chaetal tips). Scale bars: A: 2.7 mm, B: 1 mm, C: 0.2 mm, D: 0.5 mm, E: 0.3 mm, F: 50 µm, G: 30 µm.

FIGURE 8. Bradabyssa indica n. sp., holotype (MNHN IA-Type 1797). A. Dorsal view. B. Ventral view. C. Anterior end, chaetigers 1 – 3, dorsal view. D. Chaetiger 1, basal notochaetal region. E. Chaetigers 2 – 3, left parapodial neurochaetae, ventral view. F. Chaetiger 4, left parapodial neurochaetae, ventral view. Scale bars: A: 0.4 mm, B: 0.3 mm, C: 0.1 mm, D – F: 40 µm.

FIGURE 11. Bradabyssa sachalina (Annenkova-Chlopina, 1922) n. comb. A. Holotype (ZIRAS 26659), right lateral view, tunic eroded. B. Same, anterior end, ventral view (arrow points to gonopodial lobe). C. Non-type specimen (CAS 19105), right lateral view, tunic complete. D. Same, head exposed, frontal view, right palp missing (Br: branchial scars, LL: lateral lip, VPO: ventral pharyngeal organ). E. Non-type specimen (ZIRAS 26661), chaetiger 13, right parapodium, notochaetae (insets: basal, medial and distal regions). F. Same, neurochaetae. Scale bars: A: 3.6 mm, B: 0.8 mm, C: 3.9 mm, D: 0.2 mm, E: 100 µm, F: 120 µm.

FIGURE 26. Bradabyssa mammillata (Grube, 1877) n. comb. A. Syntypes (ZMB 689), oblique dorsal right lateral view. B. Smaller syntype (ZMB 689), anterior end, dorsal view. C. Same, anterior end, ventral view (arrows point to gonopodial lobes). D. Non-type specimen (MNHN 186 a), head, palps and many branchiae removed (Br: branchial scars, LL: lateral lip, NL: nephridial lobe, PS: palp scar). E. Smaller syntype (ZMB 689), chaetiger 5, right parapodium, notochaetae (insets: basal, medial and distal chaetal regions). F. Syntype (ZMB 688), chaetiger 8, right parapodium, neurochaetae. G. Non-type specimen (MNHN 354), juvenile, chaetiger 19, right parapodium, neurochaetae. Scale bars: A: 3.3 mm, B: 2.3 mm, C: 1.4 mm, D: 1.0 mm, E, F: 0.4 mm, G: 130 µm.

FIGURE 33. Bradabyssa hartmanae n. sp.: A. Holotype (SIORAS 866), dorsal view. B. Same, anterior end, dorsal view. C. Paratype (ECOSUR 185), prostomium with branchiae and palps removed, frontal view (1 – 4: branchial scars, PS: palp scar). D. Holotype, anterior end, ventral view. E. Paratype (ECOSUR 185), chaetiger 6, left parapodium, anterior view (No: distal notochaetal region, Ne: neurochaetal tip). Scale bars: A: 1 mm, B: 0.25 mm, C: 0.1 mm, D: 0.27 mm, E: 150 µm.

FIGURE 34. Bradabyssa ilyvestis (Hartman, 1960) n. comb.: A. Holotype (LACM 538), dorsal view, after a few seconds immersion in methyl green stain. Paratype (LACM 539), B. Median chaetiger, right parapodium, anterior view. C. Same, notochaetal basal, medial and distal regions. D. Same, neurochaetal basal, medial and distal regions. Scale bars. A: 0.9 mm, B: 0.1 mm, C – D: 30 µm.

FIGURE 35. Bradabyssa intoshi (Caullery, 1944) n. comb., holotype (ZMA 1600). A. Anterior end, dorsal view. B. Anterior end, ventral view (arrow points to base of left gonopodial papillae). C. Chaetiger 13, right parapodium (inset: neurochaetal tips). Scale bars. A: 0.6 mm. B: 0.5 mm, C: 50 µm.

FIGURE 36. Bradabyssa jirkovi n. sp., holotype (SIORAS 6085). A. Dorsal view. B. Anterior end, dorsal view. C. Same, ventral view (arrow points to gonopodial lobes, neurochaetae removed). Paratypes (ECOSUR 186): D. Head, frontal view, palps removed, branchiae on site. E. Same, close-up, right neuropodia from chaetigers 4 – 7 (arrow points to gonopodial lobe, most papillae brushed off). F. Chaetiger 14, right parapodium, notochaetal basal, subdistal and distal regions. G. Same, neurochaetal basal, medial and distal regions. Scale bars. A: 1.5 mm, B: 0.9 mm, C: 0.6 mm, D: 0.3 mm, E: 0.2 mm, F: 40 µm, G: 80 µm.

FIGURE 37. Bradabyssa kirkegaardi n. sp., holotype (ZMUC 1781). A. Dorsal view. B. Ventral view. C. Anterior region, ventral view. Non-type specimen (SMF Sonne 136): D. Anterior chaetiger, right parapodium, anterior view. E. Same, lower notochaetae, basal and medial regions. F. Same, neurochaetal basal, medial and distal regions. Scale bars. A: 0.7 mm, B: 0.8 mm, C: 0.4 mm, D: 0.1 mm, E: 15 µm, F: 30 µm.

FIGURE 38. Bradabyssa mezianei n. sp., holotype (MNHN IA-Type 1793). A. Dorsal view. B. Ventral view. Paratype (MNHN IA-Type 1794): C. Midbody chaetiger, right parapodium, anterior view. D. Same, basal, medial and distal notochaetal regions. E. Same, basal, medial and distal neurochaetal regions. Scale bars. A – B: 0.4 mm, C: 0.2 mm, D – E: 20 µm.