Brada granulosa Hansen, 1880

Salazar-Vallejo, Sergio I., 2017, Revision of Brada Stimpson, 1853, and Bradabyssa Hartman, 1967 (Annelida, Flabelligeridae), Zootaxa 4343 (1), pp. 1-98 : 13-14

Brada granulosa Hansen, 1880

Figure 4

Brada granulosa Hansen, 1880: 229 –230, Pl. 5, Figs 4–6 .— Hansen 1882: 39, Pl. 7, Figs 21–22 .— Fauvel 1914: 238, Pl. 21, Figs 10–12 .— Støp-Bowitz 1948a: 47 –50, Fig. 13 (syn.).— Støp-Bowitz 1948b: 46 –47, map.— Jirkov & Filippova 2001: 352, Figs 1–4 .— Oug et al. 2011: 15, unnumb. figs a–c (repeated from Støp-Bowitz, 1948a).— Moore 1909a: 143. (non Malmgren, 1867).

Brada normani McIntosh, 1908: 543 –544, Pl. 12, Figs 4 , 5 , Pl. 12A, Fig. 10 .

Type material. Northwestern European Arctic Ocean. Four syntypes ( ZMUB 2196), partially dehydrated, damaged, Norwegian North Atlantic Expedition, Sta. 275 (74°08' N, 31°12' E), 269 m, greenish clay, 1877 (26–35 mm long, 7–9 mm wide, cephalic cage 1 mm long (only one neurochaetae left in one specimen), 23–25 chaetigers). Four complete syntypes ( ZMUB 2197), and fragments of 3 others, Norwegian North Atlantic Expedition, in two stations: 337 (76°23' N, 16°43' E), off southernmost point of Spitzbergen, Norway, no depth data, and 370 (78°48' N, 08°37' E), off the westernmost point of Spitzbergen, Norway, about 915 m, 1877 (largest used for redescription, others 22–31 mm long, 9.5–12 mm wide, cephalic cage 0.5–1.0 mm long, 22–23 chaetigers; one anterior fragment had ripe ovaries with eggs about 200 µm).

Additional material. Five specimens ( ZMUB 26383), Voringen, Sta. 267 (no further data) (13–41 mm long, 4.5–9.0 mm wide, cephalic cage 0.5 mm long, 22–24 chaetigers). Two specimens ( ANSP 2853), off Fish Island, outside of Hebron, Labrador, Canada, 137 m, mud, 25 Aug. 1908, O. Bryant, coll. (38–44 mm long, 10 mm wide, cephalic cage 0.9–1.0 mm long, 24 chaetigers; gonopodial lobe in chaetiger 4).

Description. Largest syntype (ZMUB 2197) complete, brownish, slightly damaged ( Fig. 4A ). Body fusiform, blunt at both ends; 36 mm long, 12 mm wide, cephalic cage 1 mm long, 21 chaetigers. Papillae long, conical, mucronate, mucron capitate, larger dorsally, shorter ventrally; arranged in 6–7 alternating transverse series dorsally, 4–5 ventrally; all papillae with large sediment particles, forming large tubercles in first few chaetigers, particularly close to chaetal lobes (more visible in larger syntypes).

Anterior end observed by dissection in an anterior fragment (ZMUB 2197). Prostomium low, dark-gray, with some black spots ( Fig. 4C ). Eyes not seen. Caruncle wide basally, extended posteriorly, not reaching branchial plate margin. Palps dark gray; palp keels rounded, slightly darker. Lips darker; lateral lips thick, well developed; ventral lip reduced. Dorsal lip darker, triangular, clearly separated from lateral lips.

Branchiae thick cirriform, sessile on branchial plate, arranged in a single marginal row, filaments separated into two lateral groups, each with 4 filaments. Largest branchiae as long as, and slightly thinner than palps. Nephridial lobes on branchial plate not seen.

Cephalic cage chaetae as long as 1/36 of body length, or 1/12 as long as body width. Chaetiger 1 with 1–2 thin chaetae per ramus, arranged in short lateral series.

Anterior dorsal margin of first chaetiger triangular, papillated. Anterior chaetigers without especially long papillae; in larger specimens (ANSP 2853) with fused tubercles forming a sandy crust. Chaetigers 1–3 gradually decrease in size. Chaetal transition from cephalic cage to body chaetae abrupt, with curved anchylose neurospines from chaetiger 2. Gonopodial lobes present in chaetiger 4, short, digitate, grayish ( Fig. 4B ).

Parapodia well developed, lateral; median neuropodia ventrolateral ( Fig. 4D ). Notopodia and neuropodia close to each other. Notopodia digitate thin lobes, covered by thin layer of sediment particles, surrounded by larger papillae. Neuropodia larger, truncate conical lobes, covered by thin layer of sediment, with about 3 pre- and 2 postchaetal papillae, much larger than neurochaetal lobe.

All notochaetae multiarticulate capillaries; median notochaetae arranged in short series, 2 chaetae per ramus, one with short articles basally, becoming longer medially and even longer distally, the other with very long articles, increasing in length distally ( Fig. 4E ). Notochaetae as long as 1/5–1/6 body width. Neurochaetae multiarticulate capillaries in chaetiger 1. Falcate, subdistally expanded blunt neurospines from chaetiger 2, arranged in oblique series, 5–6 per bundle, tips falcate ( Fig. 4F ), more pronounced in median and posterior chaetigers ( Fig. 4G ). Tips appearing hooded ( Fig. 4G , inset).

Posterior end globose, pygidium with anus terminal, as a vertical slit; anal cirri absent.

Variation. Adult specimens 13–44 mm long, 9.5–12.0 mm wide, 22–24 chaetigers.

Remarks. Brada granulosa Hansen, 1880 and B. granosa Stimpson, 1853 are very similar because, in both, dorsolateral papillae are fused forming transverse series of sandy tubercles. However, they differ by the relative shape of neurochaetae and pattern of ventral papillae: in B. granulosa neurochaetae are more falcate, with wide, blunt tips, and ventral papillae are of a similar size, whereas in B. granosa neurochaetae are less falcate, with tapered tips, and ventral papillae alternate in size.

Also, B. normani McIntosh, 1908 was regarded as a junior synonym of B. inhabilis ( Rathke, 1843) by Haase (1915: 177, 206), and by Støp-Bowitz (1948a:40), who also included McIntosh (1908) in Remarks but failed to list B. normani in the synonymy. Despite McIntosh’s (1908) short description, the illustrations support the case for B. normani being regarded as a junior synonym of B. granulosa because the dorsal papillae are long and tapered, and neurochaetal tips are wide. Herein B. normani is regarded as a junior synonym of B. granulosa.

Distribution. Arctic and subarctic environments, reaching about 1000 m depth.

Figures

FIGURE 1. Morphological features of Brada Stimpson, 1853, and Bradabyssa Hartman, 1967. A. Brada granulosa Hansen, 1880, lateral view. B. neurochaeta. C. Bradabyssa (olim Trophonia) rugosa (Hansen, 1880) n. comb., lateral view. D. Median parapodium. E. Notochaeta. F. Neurochaeta (modif. after Hansen 1880).

FIGURE 2. Brada granosa Stimpson, 1853. A. Neotype (USNM 1422407), lateral view, anterior end to the right (inset: small paratype, lower right in Fig. 3 A). B. Same, anterior end, ventral view (arrows point to gonopodial lobes). C. Non-type specimen (USNM 22851), head dissected, branchiae and palps removed (Br: branchial scar, PS: palp scar). Scale bars: A: 1.7 mm, B: 1.3 mm, C: 0.4 mm.

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 5. Brada incrustata Støp-Bowitz, 1948. A. Lectotype (ZMUB 18605), right lateral view, anterior end above. B.

FIGURE 6. Brada inhabilis (Rathke, 1843), smaller, non-type specimen (ZMUB 34537). A. Right lateral view. B. Anterior end, lateral view (arrow points to gonopodial papilla). C. Head, frontal view (palps and branchiae removed; Br: branchial scar, PS: palp scar). D. Median parapodium (greenish after methyl-green staining). E. Anterior parapodium notochaetae (inset: enlargement of notochaeta). F. Posterior parapodium, two neurochaetae (insets: neurochaetal tips). Scale bars: A: 2.1 mm, B: 1.3 mm, C: 0.4 mm, D: 0.3 mm, E: 75 µm, F: 120 µm.

FIGURE 10. Bradabyssa minuta (Amoureux, 1986) n. comb., syntypes (MNHN 834). A. Five syntypes, mostly without globose foraminiferans. B. Three syntypes with globose foraminiferans on tunic. C. Median chaetiger, right parapodium (insets: notochaetal basal regions, neurochaetal tips). Scale bars: A: 0.4 mm, B: 0.3 mm, C: 65 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 12. Bradabyssa alaskensis n. sp. A. Holotype (LACM 2598), right lateral view, tunic slightly eroded. B. Same, anterior end, lateral view (palp and branchiae slightly exposed). C. Paratype (LACM 2599), head, frontal view, palps and branchiae removed (Br: branchial scars, PS: palp scar). D. Holotype (LACM 2598), anterior end, ventral view (arrows point to everted gonopodial lobes). E. Same, close-up of left gonopodial lobe. F. Same, chaetiger 10, right parapodium, frontal view. G. Paratype (LACM 2599), chaetiger 5, right parapodium, notochaetae (insets: medial and distal regions). H. Same, neurochaetae. I. Same, chaetiger 23, right parapodium, neurochaetae. Scale bars: A: 2.5 mm, B: 0.9 mm, C: 0.6 mm, D: 1.2 mm, E: 0.3 mm, F: 0.5 mm, G: 50 µm, H: 100 µm, I: 90 µm.

FIGURE 13. Bradabyssa antarctica (Hartman, 1978) n. comb., holotype (USNM 46823). A. Dorsal view. B. Anterior end, dorsal view. C. Posterior end, dorsal view (methyl green staining). D. Chaetiger 7, right parapodium, posterior view. E. Same, neurochaetae. Scale bars: A: 0.7 mm, B, C: 0.2 mm, D: 60 µm, E: 40 µm.

FIGURE 21. Bradabyssa elinae n. sp. A. Holotype (SMNH 55750), oblique, left lateral view. B. Anterior end, dorsal view. C. Anterior end, ventral view (arrows point to gonopodial lobes). D. Paratype (SMNH 55751), chaetiger 16, left parapodium, anterior view. E. Same, notochaetal basal, medial and distal regions. F. Same, neurochaetae (inset: chaetal tip, arista twisted). G. Holotype (SMNH 55750), posterior end, dorsal view. Scale bars: A: 1.5 mm, B: 0.6 mm, C: 0.7 mm, D: 150 µm, E: 70 µm, F: 130 µm, G: 0.5 mm.

FIGURE 22. Bradabyssa grangieri n. sp. A. Holotype (USNM 1422382), oblique dorsolateral view, most tunic removed. B. Anterior end, oblique dorsal view. C. Anterior end, ventral view (arrows point to gonopodial lobes). D. Posterior end, oblique dorsal view. E. Paratype (USNM 1422383), head, frontal view, palps and branchiae removed (Br: branchial scars, PS: palp scar). F. Same, chaetiger 10, right parapodium, basal, medial and distal notochaetal regions. G. Same, aristate neurospines, one with a subdistal constriction. H. Non-type specimen (USNM 1422385), chaetiger 10, right parapodium, typical neurospine tips. Scale bars: A: 5.3 mm, B: 1.1 mm, C: 1 mm, D: 1.2 mm, E: 0.9 mm, F: 60 µm, G, H: 120 µm.