Flabelligera affinis Sars, 1829

Salazar-Vallejo, Sergio I., 2012, Revision of Flabelligera Sars, 1829 (Polychaeta: Flabelligeridae) 3203, Zootaxa 3203 (1), pp. 1-64 : 10-12

Flabelligera affinis Sars, 1829

Figure 1

Flabelligera affinis Sars 1829:31–34 , Pl. 3, Figs. 16–19 ; Cunningham & Ramage 1888:677–678, Pl. 46, Fig. 40A–B; de Saint- Joseph, 1894:96–101, Pl. 5, Figs. 121–123 (partim); Newbigin 1900:190 (partim); Moore 1909b:143; Gunther 1912:11– 12; Haase 1915:183–186, Pls. 1–2, Textfig. 1; McIntosh 1915:107–114, Pl. 89, Fig. 2 , Pl. 96, Fig. 10 , Pl. 104, Figs. 7–7b (syn.); Chamberlin 1919b:398–399; Fauvel 1927:113, Figs. 40a–f; Monro 1930:160, Fig. 63; Rioja-LoBianco 1931:88– 90, Pl. 24, Figs. 1–3 ; Monro 1936:164; Støp-Bowitz 1948a:8–13, Fig. 1 & map; Støp-Bowitz 1948b:30–33, map; Wesen- berg-Lund 1950:35; Pettibone 1954:289–290, Fig. 33e–g (partim); Pettibone 1956:563; Hartmann-Schröder 1971:370– 372, Fig. 130; Amouroux 1974:179–180; Hartmann-Schröder 1996:417–419, Fig. 203; Jirkov & Filippova 2001:360–361, Figs. 1–7o ; Oug et al. 2011:6, Figs. a, b.

Chloraema dujardini de Quatrefages 1849:282–285 , Pl. 9, Figs. 1–2 , Pl. 10, Figs. 1–5 .

Chloraema pellucidum Sars 1869:253 nomen nudum.

Chloraema pellucidum Sars in Sars 1872:409; 1873:252–261, Pl. 16, Figs. 9–20 .

Type material. Northwestern France. Syntypes of Chloraema dujardini de Quatrefages, 1849 ( MNHN-465 ), damaged, St. Vaast (Saint-Vaast-la-Hogue, 49º35'19" N, 01º15'58" W), Normandy .

Additional material. Norway. Two specimens ( ZMUB-2253 ), Tromsø, no further data, Danielsen, coll. (largest specimen used for description). Two specimens ( ZMUB-2263 ), damaged, Holmer Graa, on corals, Friele & Hansen, coll. Three specimens ( ZMUB-18636 ), heavily contracted, tunic transparent, Tfjinsgaarden, Stat. 40, App., coll. An anterior fragment ( ZMH-23598 ), Liefdefjorden, Spitzbergen, SPE-92/68, 7 Aug. 1992, Fucusstation, Taucherprobe, Plante & Wagner, coll . Greenland. Several specimens ( USNM-148624 ), Walrus Grounds, Murchison Sound , NW Greenland (77°45' N), first haul otter trawl, 45 fathoms, 7 Aug. 1938, number 127, R.A. Bartlett coll. (some specimens without tunic; one with body broken by pharynx contraction, has about 20 branchial filaments). Several specimens ( USNM-148624 ), Walrus Grounds, Murchison Sound, NW Greenland (77°38' N), otter trawl, 60 fathoms, 8 Aug. 1938, number 144, R.A. Bartlett coll. One mature female ( USNM-165293 ), residue of dredge, see oil tin No. 1, 9 Aug. 1943, number 3, R.A. Bartlett, coll. (22 chaetigers, no cephalic cage; 2–3 transverse rows of clavate papillae) . Iceland. One specimen ( MNHN-184 a), most tunic removed, Patieksfjorde, R/ V Pourquoi-pas?, Stat. 10 bis, 5 m, 22 Aug. 1929 (60 mm long, 5.5 mm wide, cephalic cage 2.5 mm long, 58 chaetigers). One specimen ( MNHN-184 b), R/ V La Manche Expedition, 1892 (no further data) (noto- and neuropodial lobes distinct) . Ireland. One specimen ( MNHN-457 ), mature female, Clew Bay, West Ireland , no more data, P. Fauvel, coll . France. One specimen ( MNHN-457 b), St. Anne, Cherbourg , Aug. 1916, P. Fauvel, coll. An anterior fragment ( MNHN-457 c), Urville , Regnéville-sur-Mer , Basse Normandie, 18 Jul. 1913, P. Fauvel, coll. One specimen ( MNHN-457 d), Cherbourg , 19 Apr. 1924, P. Fauvel, coll. (tunic with two large nematodes and a large rounded vesicle, probably made by another type of parasite). Three specimens ( MNHN-507 a), brownish, compressed, many neurochaetae broken, Tathiou, Apr. 1895, no further data. One specimen ( MNHN-507 ), without posterior region, anterior end exposed, distorted, eroded, Urville (Urville-Nacqueville, 49º40'29” N, 01º44'15” W), 18 Jul. 1913, P. Fauvel, coll. One specimen ( MNHN-507 b), splendid, St. Anne , Cherbourg, Aug. 1916, P. Fauvel, coll. One specimen ( MNHN-507 c), twisted, partially dehydrated, anterior end exposed, Cherbourg, 19 Apr. 1924, P. Fauvel, coll. (large nematode emerging from the tunic dorsally, and a globular blister or cyst, ventrally, in chaetiger 11). Arctic Canada. One specimen ( USNM-11181 ), gift by Chas Lütken, ZMUC. Several specimens ( USNM-246975 ), damaged, Stat. 720 (69°21.5' N, 81°40.5' W), B21/P72, Fore Basin, 8–9 m, 10 Oct. 1955, Grainger , coll. (two specimens with two different nematodes in the tunic, one black and white, the other pink) .

Description. Body pale (ZMUB-2253), tapering posteriorly ( Fig. 1A, E ); 57 mm long, 7 mm wide, cephalic cage 5 mm long, 52 chaetigers. Tunic thick, transparent, densely papillated, somehow damaged, with cover of fine sediment particles; papillae long, clavate, mucronate.

Cephalic hood not exposed. Prostomium elevated, rounded, with four dark, sometimes faded eyes. Caruncle well developed, wide, exceeding the branchial plate margin, distally truncate ( Fig. 1D ). Palps pale, thick, twice as long as branchiae, about four times thicker than branchiae, often exposed ventrally ( Fig. 1C ); palp bases rounded, elevated. Lateral lips well developed, fused to dorsal lip; ventral lip triangular, without ventral projection.

Branchiae in two lateral groups, each with filaments in concentric rows, each group with about 60 filaments. Longest branchiae about half as long as palps. Nephridial lobes paired, lateral to caruncle, about the median distance of branchial groups ( Fig. 1D ).

Cephalic cage chaetae about 1/5–1/9 body length, slightly shorter than body width ( Fig. 1B ). Chaetiger 1 involved in cephalic cage, with about 34 noto- and 20 neurochaetae per side. Anterior dorsal margin of chaetiger 1 papillated. Anterior chaetigers without especially long papillae. Chaetiger 1 longer than chaetigers 2–3. Chaetal transition from cephalic cage to body chaetae abrupt, chaetiger 2 with multiarticulated neurohooks. Gonopodial lobes present in chaetigers 5 and 6.

Parapodia well developed, lateral; median neuropodia ventrolateral. Notopodial lobes projecting, basally rounded, distally depressed, truncate, without different papillae. Neuropodial lobes projected, digitate, without different papillae. Noto- and neuropodia widely separated.

Median notochaetae arranged in short transverse rows; all notochaetae multiarticulated capillaries, as long as 1/3 body width,7–8 per bundle, articles best developed distally, very short throughout all the chaetae ( Fig. 1F ), some chaetae with articles ill-defined. Neurochaetae multiarticulated capillaries in chaetiger 1. Multiarticulated neurohooks from chaetiger 2 ( Fig. 1G ), one per ramus (most with a replacement hook). Handle articulation medially placed, articles irregular (Fibonacci sequence); three articles: small, medium-sized and long. Other articles anchylosed, small, continued to bending region. Crest slightly wider than handle, yellowish in anterior chaetigers, with darker distal half in median and posterior chaetigers; width:length ratio 1:4.

Posterior end truncated; pygidium with anus terminal, not covered by tunic, thick muscular ring, without anal cirri.

Variation. Two adult specimens of very different sizes (ZMUB-2253) show that the cephalic cage is sizedependent. The number of chaetigers was the same (52) but one specimen was three times longer than the other (22 mm vs 57 mm). The articulation of the neurohook handle is a consistent feature. The specimens have 2–3 distinct, larger articles and although the syntypes are very small, and the number of articles might be regarded as sizedependent, the number of large articles is not.

Remarks. Flabelligera affinis Sars, 1829 resembles F. orensanzi n. sp. They differ because in F. affinis the tunic does not include sediment particles and the neurohooks have pale, tapering crests, whereas in F. orensanzi n. sp. the tunic often has a slight sediment cover and neurohooks crests are medially widened and dark red.

The original description of F. affinis by Sars (1829) indicated his materials were from rocky and muddy bottoms and that it had a leathery dirty-green tunic. Rathke (1843) described Siphonostoma vaginiferum from muddy bottoms, and indicated it has an opaque, rugose tunic. Rathke’s approach is herein followed to restrict F. affinis to the Flabelligera specimens with a transparent tunic, whereas those having an opaque tunic are regarded as belonging to a distinct species, F. vaginifera (see below). Chloraema pellucidum Sars in Sars, 1872, was regarded as a junior synonym of F. affinis by Haase (1915) and later authors. However, it was originally recognized as an independent species by Michael Sars, who had also described F. affinis . Because F. affinis has been restricted to include specimens with a mostly transparent tunic, C. pellucidum is regarded as conspecific. The syntypes of C. dujardini (MNHN-465) are damaged juveniles, one broken into two pieces, and many neurohooks are broken in both specimens. Their bodies are long, fusiform, compressed, tapered posteriorly; the tunics are damaged, but without sediment particles, transparent, leaving chaetal tips uncovered. They are 8.5–10.0 mm long, 1 mm wide, cephalic cage 1 mm long, 29–33 chaetigers. Newbigin (1900:190, 195) concluded that F. affinis includes F. dujardini , F. diplochaitus (Otto) , and F. claparedi (de Saint-Joseph) . He was correct for the first species only because F. diplochaitus is herein regarded as a distinct species, whereas F. claparedei has been transferred to Flabelliderma Hartman, 1969 ( Salazar-Vallejo 2007).

Distribution. As currently recognized, F. affinis is distributed from Arctic to cold and temperate, boreal localities. It has been recorded from many widespread localities; such records by Fauvel & Rullier (1957:376, 1959a:180, 1959b:958) for Western Africa, as well as those for Southern Africa by Day (1961:505, Fig. 8a ), or by Monro (1933:1056–1057, Text fig. 5) for Panama, are all questionable. Likewise, the records by Chamberlin (1920:21) for the Canadian Arctic, or those by Ushakov (1955:306/ 1965:283, Fig. 113A–D), and by Hobson & Banse (1981:58, Fig. 11e ), for the Northern Pacific Ocean, belong elsewhere.

Figures

FIGURE 1. Flabelligera affinis Sars, 1829. A. Non-type specimens (ZMUB-18636), dorsal view. B. Non-type specimen (MNHN- 184b), anterior end, dorsal view. C. Same, anterior end, ventral view. D. Non-type specimen (MNHN-184a), anterior end, most tunic removed, head exposed (Ca: caruncle, LL: lateral lip, NL: nephridial lobe, PL: palp lobe). E. Non-type specimen (ZMUB- 2253), lateral view, most tunic removed. F. Same, chaetiger 10, notochaetal basal and medial regions (insert: subdistal regions). G. Same, chaetiger 10, neurochaetae (inserts: articulated, median region and crest). Scale bars: A: 1.3 mm, B: 2.3 mm, C: 2 mm, D: 0.5 mm, E: 10 mm, F: 195 µm, G: 120 µm.

FIGURE 2. Flabelligera bicolor (Schmarda, 1861). A. Syntype (BMNH-1908.5.9.33), lateral view. B. Same, anterior end, lateral view. C. Another syntype (BMNH-1908.5.9.33), head, frontal view (BS: branchial scars, Ca: caruncle, LL: lateral lip, VL: ventral lip). D. Same, chaetiger 10, basal and medial notochaetal regions. E. Same, neurohook. F. Same, posterior chaetiger, neurohook (insert: handle articulation). Scale bars: A: 2.2 mm, B: 2.3 mm, C: 1 mm, D: 66 µm, E: 0.30 mm, F: 0.38 mm.

FIGURE 3. Flabelligera bophortica Annenkova-Chlopina, 1924, syntypes (ZIRAS-27043). A. Best syntype, lateral view. B. Same, anterior end, oblique, ventrolateral view. C. Same, head, frontal view (BS: branchial scars, Ca: caruncle, DL: dorsal lip, LL: lateral lip, PS: palp scar, VL: ventral lip). D. Same, neuropodia 4–7, oblique, ventral view showing gonopodial lobes (GL), E. Same, chaetiger 10, notochaetae (inserts: basal, medial and subdistal regions). F. Same, posterior chaetiger, neurohooks (inserts: handle articulation and crest close-up). Scale bars: A: 3.3 mm, B: 1.3 mm, C: 0.5 mm, D: 0.8 mm, E: 320 µm, F: 260 µm.

FIGURE 7. Flabelligera luctator (Stimpson, 1856), topotype specimens (BMNH-1936.10.16.152–7). A. Complete specimens. B. Largest syntype, anterior end, dorsal view. C. Same, head, frontal view (LL: lateral lip, NL: nephridial lobe, PS: palp scar). D. Same, chaetigers 4–7, ventral view (GL: gonopodial lobes). E. Same, chaetiger 5, neurohooks (insert: articulated portion of handle). F. Same, chaetiger 11, neurohook (insert: articulated portion of handle). Scale bars: A: 10 mm, B: 2.3 mm, C: 0.5 mm, D: 0.8 mm, E: 225 µm, F: 160 µm.

FIGURE 8. Flabelligera multipapillata Hartmann-Schröder, 1965, Holotype (ZMH-14911). A. Dorsal view. B. Anterior end, dorsal view; greenish due to methyl green staining. C. Same, chaetiger 12, neurohook (insert: basal and distal regions, enlarged). Scale bars: A: 1.7 mm, B: 0.6 mm, C: 150 µm.

FIGURE 9. Flabelligera nuniezi n. sp. A. Holotype (MNHN/1530), dorsal view. B. Same, ventral view. C. Same, chaetigers 3– 9, left neuropodia, showing gonopodial lobes by arrows. D. Same, chaetiger 10, neurohook. E. Same, chaetiger 22, neurohook. F. Non-type specimen (ZMUC unnumb.), median neurohook (insert: close up of handle anchylosed region). Scale bars: A: 1.3 mm, B: 1.1 mm, C: 0.2 mm, D: 90 µm, E: 100 µm, F: 250 µm.

FIGURE 10. Flabelligera orensanzi n. sp. A. Holotype (MACN-In 39270), dorsal view. B. Same, anterior end, ventral view. C. Paratype (CENPAT), head, frontal view, palps and branchiae removed, prostomium slightly eroded (BS: branchial scars, LL: lateral lip, PS: palp scar). D. Paratype (ECOSUR), chaetiger 9, neurohook (insert: articulated region of handle). E. Non-type specimen (MNHN-884), head, frontal view, two copepod parasites indicated by arrows (Ca: caruncle, BS: branchial scar, VL: ventral lip). Scale bars: A: 0.8 mm, B: 1 mm, C: 0.5 mm, D: 225 µm, E: 0.25 mm.

FIGURE 11. Flabelligera pergamentacea Ehlers, 1913. A. Non-type specimen (ZIRAS-16343), lateral view. B. Same, anterior end, dorsal view. C. Large syntype (ZMB-6774), broken into two pieces, dried out. D. Non-type specimen (ZMH-8592), dried out. E. Non-type specimen (ZIRAS-16343), agglutinated sediment particles on tunic surface. F. Same, chaetiger 10, neurohook (inserts: crest and handle articulated region). Scale bars: A: 2.5 mm, B: 0.5 mm, C: 1.3 mm, D: 1.5 mm, E: 35 µm, F: 50 µm.

FIGURE 12. Flabelligera picta Ehlers, 1913. A. Non-type specimen (ZMH-24295), dorsal view. B. Same, anterior end, dorsal view. C. Same, anterior end, ventral view. D. Same, median chaetiger, notochaetae (inserts: basal and subdistal chaetal regions). E. Same, chaetiger 5, neurohooks (inserts: articulated regions and crests). F. Paratype (ZMH-8591), lateral view. Scale bars: A: 3 mm, B: 0.8 mm, C: 0.7 mm, D: 250 µm, E: 260 µm, F: 1 mm.

FIGURE 13. Flabelligera salazarae n. sp. A. Holotype (ECOSUR 117), dorsal view. B. Same, anterior end, ventral view. C. Paratype (ECOSUR), head, frontal view, palps and most branchiae removed (BS: branchial scars, Ca: caruncle, LL: lateral lip, NL: nephridial lobe, PS: palp scar, VL: ventral lip). D. Holotype, chaetigers 3–6, left side, ventral view, methyl green stained, arrows point gonopodial lobes. E. Same, chaetiger 24, neurohook (inserts: multiarticulated region and crest). Scale bars: A: 1.5 mm, B: 1 mm, C: 0.2 mm, D: 0.4 mm, E: 230 µm.

FIGURE 14. Flabelligera sordida (de Quatrefages, 1849) reinstated. A. Holotype (MNHN-466), dorsal view. B. Non-type specimen (BMNH-1955.8.22.1/3), anterior end, dorsal view. C. Same, posterior end, dorsal view. D. Non-type specimen (MNHN-184), irregular, lateral view. E. Same, median chaetigers showing globular papillae. F. Same, head, frontal view, methyl green staining. G. Non-type specimen (BMNH-1955.8.22.1/3), chaetiger 4, notopodium (insert: globular papillae from posterior chaetiger). H. Same, chaetiger 12, neurohook (insert: multiarticulated region). Scale bars: A: 1.5 mm, B: 1.1 mm, C: 1.3 mm, D: 3.3 mm, E: 0.4 mm, F: 0.2 mm, G: 0.6 mm, H: 320 µm.

FIGURE 15. Flabelligera vaginifera (Rathke, 1843). A. Non-type specimen (ZMUB-27502), dorsal view. B. Same, some tunic fragments removed, anterior end, dorsal view. C. Non-type specimen (ZMUB-2254), head compressed by cover slip, branchial plate dislodged (BS: branchial scars, LL: lateral lip, PS: palp scar). D. Another specimen (ZMUB-2273), anterior end, ventral view (arrows indicate gonopodial lobes). E. Same, right chaetigers 5–6 (arrows indicate gonopodial lobes). F. Same, chaetiger 9, neurohooks (insert: multiarticulated region). Scale bars: A: 2 mm, B: 2.2 mm, C: 0.3 mm, D: 3 mm, E: 1.2 mm, F: 0.45 mm.

FIGURE 16. Annenkova mastigophora (Annenkova, 1952) n. comb. A. Lectotype (ZIRAS-27061), oblique, lateral view. B. Same, chaetigers 4–7, left neuropodia, asterisks on gonopodial lobes. C. Same, chaetiger 10, basal, medial and distal notochaetal regions. D. Same, chaetigers 7–11, neuropodia. E. Same, falciger with incurved crest tip (insert: complete neurohook). F. Same, falciger with perpendicular crest tip (insert: complete neurohook). G. Same, falciger with broken crest tip. Scale bars: A: 5 m, B: 0.2 mm, C: 40 µm, D: 1 mm, E: 50 µm, F: 57 µm, G: 50 µm.

FIGURE 17. Annenkova mastigophora (Annenkova, 1952) n. comb. A. Non-type specimen (CAS-26832), dorsal view. B. Same, ventral view. C. Same, head, palps and most antennae removed (BS: branchial scars, Ca: caruncle, LL: lateral lip, PS: palp scar). D. Same, chaetiger 2, neurohook (insert: multiarticulated handle region). E. Same, chaetiger 15, neurohooks (inserts: oligoarticulated regions and crests). F. Same, chaetiger 25, right parapodium, posterior view (notice the single articulation). Scale bars: A: 1.2 mm, B: 1.1 mm, C: 0.25 mm, D: 140 µm, E–F: 190 µm.

FIGURE 18. Flabegraviera mundata (Gravier, 1906) n. comb. A. Syntypes (MNHN-561), larger one in lateral view, smaller one in ventral view. B. Non-type specimen (MNHN-A183), cross-section, posterior chaetigers. C. Smaller syntype, anterior end, ventral view (Ov: ova, Pa: palp). D. Non-type specimen (USNM-271533), head, frontal view, palps and branchiae removed (BS: branchial scars, LL: lateral lip, PS: palp scar). E. Smaller syntype, chaetiger 15, distal notochaetal regions (insert: articulated tip). F. Same, chaetiger 4, neurohooks (inserts: multiarticulated regions and bending portion). G. Same, chaetiger 14, neurohooks (insert: distal part of handle and crest). Scale bars: A: 5.2 mm, B: 1 mm, C: 1.3 mm, D: 0.4 mm, E: 110 µm, F: 230 µm, G: 350 µm.

FIGURE 19. Flabegraviera profunda n. sp. A. Holotype (USNM-26611), dorsal view. B. Same, anterior end, ventral view. C. Paratype (USNM-unnumb.), head, frontal view, palps and branchiae removed (BS: branchial scars, DL: dorsal lip, LL: lateral lip, PS: palp scar). D. Holotype, posterior chaetiger. E. Same, neurochaetae (insert: neurochaetal tips). F. Smaller paratype, median chaetiger, neurohooks showing poorly developed crest bending sector. Scale bars: A:5.8 mm, B: 1.2 mm, C: 0.6 mm, D: 2 mm, E: 0.4 mm, F: 75 µm.

FIGURE 20. Flabehlersia induta (Ehlers, 1897) n. comb. A. Syntype (ZMH-4875), lateral view. B. Another syntype (ZMH- 4875), dorsal view. C. Same, anterior end, oblique ventral view. D. Another syntype (SMNH unnumb.), head, frontal view, methyl green stained, palps and most branchiae removed (BS: branchial scars, Ca: caruncle, LL: lateral lip, PS: palp scar). E. Another syntype (ZMH-4875), chaetigers 5–6, arrows point to gonopodial lobes. F. Median chaetiger (anterior fragment ZMH- 4875), methyl green stained, frontal view. G. Another syntype (ZMH-4875), chaetiger 10, right parapodium. H. Same, neurohook of same. Scale bars: A: 3.6 mm, B: 3.2 mm, C: 1.6 mm, D–E: 0.3 mm, F: 1 mm, G: 350 µm, H: 190 µm.