BULLETIN OF MARINE SCIENCE, 67(1): 25–37, 2000
SPECIES OF NEREIDIDAE FROM THE CAPBRETON CANYON (BAY OF BISCAY, NORTHEAST ATLANTIC)
Jorge Núñez, Florencio Aguirrezabalaga and Argiloa Ceberio
ABSTRACT Five species of Nereididae collected from the Capbreton Canyon, southeastern part of the Bay of Biscay, in deep waters (480–1040 m) are described. Three species are new to the Atlantic: Nicon sinica Wu and Sun, 1979, Ceratocephale cf. pacifica (Hartman, 1960) and C. cf. andaman Hylleberg and Nateewathana, 1988; and one, Ceratocephale loveni Malmgren, 1867 is new record for the Iberian fauna. The distributional range for Eunereis longissima (Johnston, 1840) in the Iberian region is extended.
The Capbreton Canyon begins at 250 m from the coastline near Capbreton and stretches from the east to the west along 135 nautical mi. This canyon divides the continental shelf of the Bay of Biscay into two zones. The benthic macrofauna of this canyon was studied by Le Danois (1948), who distinguished two levels: the upper one between 70 and 500 m depth was dominated by scleractinian and alcionarian facies, whereas the lower one be- tween 500 and 1000 m was characterized by the presence of hexactinelids and echino- derms. Four oceanographic cruises were carried out on board the RV CÔTE D’AQUITAINE from 1987–1990, in which studies of macrofauna communities were included. The first results relating to polychaetous annelids were published by Rallo (1988), Rallo et al. (1993), García-Arberas and Rallo (1994), and San Martín et al. (1996). The objective of this work was to extend the knowledge of the polychaete fauna, particularly the deep-sea benthic Nereididae from the Bay of Biscay.
MATERIAL AND METHODS
The specimens belonging to the Nereididae family were collected at seven of the stations dredged during the Capbreton 1988 and 1990 cruises (Fig. 1, Table 1). Samples were made with three types of gear: a Sanders-Hessler epibenthic dredge with a mesh size of 0.5 mm, a Reineck box-corer (sample sieved with a mesh of 0.5 mm), and a Marinovitch trawl. The samples were fixed in a 10% formaldehyde-seawater solution and preserved in a 70% alcohol solution. For identification, parapo- dia from the majority of the specimens were removed and permanent slide mounts in a glycerin gel were made. The drawings were made to scale, with the aid of a drawing tube coupled to a LEICA DMLB optic microscope equipped with interference contrast optics (Nomarski). The specimens were deposited at the Museo Nacional de Ciencias Naturales de Madrid (MNCNM), Spain.
25 26 BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000
Figure 1. Map of Capbreton (Bay of Biscay) showing sampled stations.
SYSTEMATICS
Ceratocephale loveni Malmgren, 1867 (Fig. 2A–J)
Ceratocephale loveni Malmgren, 1867: 61, pl. 5, fig. 33.—Pettibone, 1963: 152, fig. 4a,b.— Hartmann-Schröder, 1971: 191.—1974: 200, figs. 19–22.—Amoureux, 1982: 42, fig. 6.—Cham- bers and Garwood, 1992: 26, figs. 40,12,24,51.—Hilbig, 1997: 297, fig. 12.2. Ceratocephale borealis Wesenberg-Lund, 1950: 18, pl. 5, figs. 19–21; pl. 6, figs. 23–26.
Material Examined.—Five spec., sta. DI-19; one spec., sta. CM-22. Description.—Six incomplete specimens without posterior end. Largest specimen with 30 setigers, 6.5 mm long and 0.4 mm wide without parapodia and setae. Preserved mate- rial brown in color with golden setae and dark brown aciculae. Anterior 11 setigers in- flated, rest of body narrow with setigers similar in width. Anterior setigers (3–11) with numerous golden setae, following setigers with fewer setae and pale golden color. NÚÑEZ ET AL.: NEREIDIDAE FROM CAPBRETON CANYON, BAY OF BISCAY 27
Table 1. Sampling data in the Capbreton Canyon during the cruise CAPBRETON 1898 −90. DI = Sanders-Hessler epibenthic dredge, CM = Marinovitch trawl, KR = Reineck box-corer.
Station Position at the beginning and the Depth (m) at the Doate N end of the tow on the sea-floor beginning and the end individuals DI 13 43° 38.36'N−20° 18.03'W 1804 017/06/8 43° 38.08'N−27° 18.14'W 100 DI 19 43° 37.48'N−12° 52.52'W 985 057/07/8 43° 37.46'N−18° 52.66'W 96 DI 21 43° 37.72'N−10° 41.83'W 588 017/07/8 43° 37.43'N−10° 41.99'W 48 DI 26 43° 42.89'N−24° 18.71'W 988 017/08/8 43° 43.25'N−29° 18.80'W 102 CM 22 43° 37.79'N−14° 42.91'W 682 017/07/8 43° 37.50'N−12° 43.09'W 65 CM 27 43° 43.19'N−24° 20.13'W 985 057/08/8 43° 44.54'N−27° 18.18'W 91 KR 24 43° 43.69'N−20° 18.03'W 906 017/07/9 Prostomium deeply incised, with pair of stout frontal antennae, as long as palps; biarticulated palps, with elongate palpostyles (Fig. 2A); eyes absent. Peristomium achaetous, short, similar in length to following segments. Four pairs of tentacular cirri similar in length, longest reaching back to setiger 3. Pharynx not everted, pair of light brown, serrated jaws visible after dissection. Parapodia of first two setigers uniramous, with triangular postsetal lobe and digitiform neuro-acicular papilla, ventral neuropodial ligule cirriform, shorter than postsetal lobe. Dorsal cirri filiform, slightly longer than inferior lobe, single ventral cirri shorter than lobe and ligule. Setae homogomph spinigers. Following parapodium biramous. Dorsal cirri on first 9 setigers with small cirrophores (Fig. 2B), parapodia from setigers 10–20 with cirrophore of dorsal cirri elongated, inflated, vascularized and ciliated (Fig. 2C). Posterior parapodia with dorsal cirri slender and very long, without elongated cirrophore. Ventral cirri bifid from setiger 3. Notopodia of setiger 7 with prominent triangular presetal ligule and short rounded postsetal lobe. Neuropodia with broad lobe and small conical neuroacicular papilla, conical presetal ligule, slightly longer, rounded postsetal lobe and blunt, triangular ventral ligule. Notopodia of setiger 10 shorter than that of setiger 7 and neuropodial lobes small. Posterior setiger (23) with parapodia smaller than those of more anterior one of the body, short notopodial lobe and elongated neuropodial lobes (Fig. 2D). Setae include homogomph and sesquigomph spinigers (Fig. 2E–F,H–J), with blade sligtlhy serrated, and 2–4 sesquigomph spinigers with strongly serrated blades (Fig. 1G). Falcigerous setae absent. Distribution.—North Atlantic: Greenland, Scandinavia, Ireland, Bay of Biscay, Nova Scotia to Virgini, Sea of Okhotsk. Pacific: Gulf of Alaska, Valparaiso, Cape Horn (50– 5000 m depth). 28 BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000
Figure 2. Ceratocephale loveni. A. anterior end, dorsal view, B. parapodium, setiger 7, C. parapodium, setiger 10, D. parapodium, posterior setiger, E. notoseta from setiger 3, F. neuroseta, shaft from setiger 3, upper bundle, G. neuroseta from setiger 3, lower bundle, H. notosetae, shafts from setiger 7, I. neuroseta from setiger 12, upper bundle, J. neuroseta, setiger 12, lower bundle. Scale: A = 0.4 mm; B–D = 237 µm; E–J = 29 µm.
Ceratocephale cf. pacifica (Hartman, 1960) (Fig. 3A–I)
Ceratocephale loveni pacifica Hartman, 1960: 94, pl. 8.—Hartman, 1968: 501. Ceratocephale pacifica.—Banse, 1977: 617, fig. 2a.—Hilbig, 1997: 299, fig. 12.2. Ceratocephale ?pacifica.—Hutchings and Reid, 1990: 82, fig. 7a–e, tables 7,9.
Material Examined.—One spec., sta. KR 24. NÚÑEZ ET AL.: NEREIDIDAE FROM CAPBRETON CANYON, BAY OF BISCAY 29
Figure 3. Ceratocephale cf. pacifica. A. anterior end, dorsal view, B. parapodium, setiger 7, C. parapodium, setiger 20, D. notosetae, shafts from setiger 7, E. neurosetae, shafts from setiger 7, upper bundle, F. neurosetae shafts from setiger 7, lower bundle, G. notosetae, shafts from setiger 10, H. neurosetae, shafts from setiger 10, I. notosetae from posterior setiger. Scale: A = 0.3 mm; B– C = 190 µm; D–I = 29 µm.
Description.— Specimen complete with 49 setigers, 4.5 mm long and 0.3 mm wide without parapodia and setae. Alcohol-preserved specimen lacks pigment patterns, pale cream in color. Anterior 3–8 setigers with robust parapodia, after that gradually decreas- ing in size. Anterior end with pale golden setae and pale brown aciculae, rest of body with transparent aciculae and setae. Prostomium wider than long, without eyes, deeply incised in front, with pair of anten- nae as long as palps; palps with elongate palpostyles (Fig. 3A). Peristomium fused to 30 BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000 prostomium, as long as setiger 3. Four pairs of short tentacular cirri, posterior pair reach- ing back to setiger 3. Pharynx everted with three dorsal papillae on oral ring, and some ventral papillae difficult to see. Jaws serrated, slightly arched, pale brown in color, with 11 teeth. Parapodia of first two setigers uniramous, small, with filiform dorsal cirri and short bifid ventral cirri. Biramous parapodia with neuropodia more robust than notopodia. Notopodial presetal ligule elongated, acicular postsetal lobe conical. Neuropodia with broad lobe and small neuroacicular crescent papilla, and ventral ligule similar in shape and size to ventral cirri. Dorsal cirri elongated, with small cirrophore on first 8 setigers (Fig. 3B), from about setiger 9 to 20 with enlarged cirrophore, wider than style, with well-developed cilia continuing dorsally forming a transverse fringe (Fig. 3C). Parapodia of median region similar except presetal ligule shorter than dorsal cirri. Ventral cirri bifid to about setiger 20. In all parapodia setae homogomph and sesquigomph spinigers, blades smooth or slightly serrated; no strongly serrated setae. Notosetae and neurosetae with long and short blades (Fig. 3D–I). Remarks.—The single small specimen collected closely resembles Ceratocephale paci- fica (Hartman, 1960), however, the enlarged cirrophores of the dorsal cirri extends from setiger 9, rather than setiger 8 as stated original description, and the bifid ventral cirri are present from setiger 1 to about 20 in our individual, whereas in the original description they are reported from setiger 1 to about 25. The specimen is therefore tentatively as- signed to C. cf. pacifica. Distribution.—Pacific: Western Mexico to central California (840–2580 m depth). Australia: Victoria, Bass Strait (1200–2350 m depth). Atlantic: Bay of Biscay (960 m depth) (this study).
Ceratocephale cf. andaman Hylleberg and Nateewathana, 1988 (Fig. 4A–I)
Ceratocephale andaman Hylleberg and Nateewathana, 1988: 4, fig. 2.
Material Examined.—One spec., sta. DI 13. Description.— One anterior fragment with 37 setigers, 7 mm long and 0.3 mm wide without parapodia and setae. Body yellowish in alcohol, lacking pigment patterns. Setigers 3–10 with robust parapodia, abundant pale golden setae, and brown aciculae. Prosto- mium deeply incised, with pair of distal antennae slightly longer than palps. Biarticulated palps with elongated palpostyles (Fig. 4A). Eyes lacking. Peristomium longer than first segment, four pairs of short tentacular cirri, longest reaching back to setigers 1–2. Phar- ynx not everted, with pair of light brown, serrated jaws. Two first pairs of parapodia uniramous, with pair of digitate parapodial lobes, dorsal cirri with short cirrophore and elongated style, and bifid ventral cirri. From setiger 3 to 30 biramous parapodia, notopodia with small acicular papillae and elongated presetal ligule; neuropodia with small acicular papillae, wide postsetal lobes, digitiform presetal ligules as long as subpodial ligules or slightly longer. Posterior setigers with neuropodial ligules reduced in size (Fig. 4F). Dorsal cirri with enlarged cirrophore from setiger 9 up to setiger 31 (Fig. 4 B–E). Ventral bifid cirri present to at least setiger 37 (end of frag- ment). NÚÑEZ ET AL.: NEREIDIDAE FROM CAPBRETON CANYON, BAY OF BISCAY 31
Figure 4. Ceratocephale cf. andaman. A. anterior end, dorsal view, B. parapodium, setiger 10, C. parapodium, setiger 11, D. parapodium, setiger 25, E. parapodium, setiger 34, F. parapodium, setiger 37, G. notosetae from setiger 10, H. neurosetae, upper bundle, I. neurosetae, lower bundle. Scale: A = 0.2 mm; B–E = 286 µm; F–H = 29 µm.
Notosetae and neurosetae homogomph spinigers with long finely serrated blades and generally two coarsely serrated neurosetae in upper bundle and two in lower bundle (Fig. 4 G–I). Aciculae slightly arched and brown in color. Remarks.— Our specimen coincide in general with the original description by Hylleberg and Nateewathana (1988), however, the cirrophores of the dorsal cirri extend from setiger 9 to 32, whereas in the original description they extended from setiger 8 or 9 to 30. The 32 BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000 main difference is the number of the setigers which have bifid ventral cirri; in the Capbreton specimen they are present from the first setiger to at least setiger 37 (the last setiger of the considered fragment), whereas the specimens of Andaman have ventral cirri only to the 26th setiger. Since these characters are variable and since we have only one specimen, we think that it is best to leave the taxon pending until we have more specimens from the area. Distribution.—Indian Ocean: Andaman Sea and Gulf of Thailand (16–68 m depth); Atlantic Ocean: Bay of Biscay (1007–1040 m depth) (this study).
Nicon sinica Wu and Sun, 1979 (Fig. 5A–L)
Nicon sinica Wu and Sun, 1979: 11, fig. 182; Wu, Sun and Yang, 1985: 60, fig. 31; Miura, 1990: 11, figs. 1–2.
Material Examined.—Four spec., sta. CM 27. Description.—Specimen complete, 34 mm long, 0.7 mm wide without parapodia and setae, for 76 setigers. Preserved specimens without pigment pattern, yellowish in color. Prostomium pyriform, slightly longer than wide. Frontal antennae frontal slender. Palps stout with globular palpostyles. Two pairs of large eyes, arranged in trapeze. Peristomium similar in length to first setiger. Tentacular cirri smooth, slender, longest one extending back to setiger 6 (Fig. 5A). First two setigers uniramous, with dorsal cirri longer than notopodial lobe, ventral cirri shorter than neuropodial lobes; cirri with similar proportions along body. In biramous parapodia notopodia with two subequal conical lobes and shorter acicular lobe, becom- ing triangular in posterior setigers; neuropodia with bluntly triangular acicular and postsetal lobes, approximately equal in length. Inferior lobe conical anteriorly and triangular pos- teriorly (Fig. 5B–E). Notosetae numerous (23–52), all homogomph spinigers with long denticulated blades (Fig. 5F). Anterior setigers with neuropodial upper bundle of spinigers (Fig. 5G,L), some parapodia also with heterogomph spinigers (parapodia 4 with 6 homogomph and 2 heterogomph setae). Setae of lower bundle all heterogomph spinigers in setiger 4 (16 setae). Remaining setigers with heterogomph falcigers in upper and lower bundles (Fig. 5H–K). Blades short and stout in posterior parapodia, with distal tooth increasingly hooked and with distinct tendon in far posterior setigers. Pygidium with two long anal cirri. Remarks.—Our specimens agree with the description of Wu and Sun (1979), Wu et al. (1985) and Miura (1990), and with revised specimens from Japanese coasts, except for the pigmentation which is present in specimens from the Asian coasts. Those from Japan are characterized by the presence of a cross and chestnut-colored spots frontally on the prostomium, at the bases of the palpophores, on the peristomium, and dorsally and ven- trally on the segments. The Capbreton Canyon specimens, appear unpigmented. We do not know if the specimens had any coloration in vivo. On the other hand, it is common that the same species presents variations in the pigmentation pattern according to the NÚÑEZ ET AL.: NEREIDIDAE FROM CAPBRETON CANYON, BAY OF BISCAY 33
Figure 5. Nicon sinica. A. anterior end, dorsal view, B. parapodium, setiger 4, C. parapodium, setiger 10, D. parapodium, setiger 23, E. parapodium, setiger 52, F. spinigerous notoseta from setiger 10, G. spinigerous neuroseta from setiger 10, H. falcigerous neuroseta, shaft, from setiger 52, I. falcigerous neuroseta, upper bundle, from setiger 52, J. falcigerous neuroseta, lower bundle, from setiger 52, K. falcigerous neuroseta, lower bundle, from setiger 23, L. spiniger neuroseta, lower bundle from setiger 23. Scale: A = 0.7 mm; B–E = 233 µm; F–L = 21 µm. 34 BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000
environment, feeding, and the stage of their biological cycle; for this reason we feel that the identification as N. sinica for Capbreton specimens is justified. Distribution.—China: Huanghai Sea, South China Sea. Japan. Atlantic: Bay of Biscay (917–954 m depth) (this study).
Eunereis longissima (Johnston, 1840) (Fig. 6A–M)
Nereis longissima Johnston, 1840: 178. Nereis (Eunereis) longissima.—Fauvel, 1923: 351, fig. 138a-d.—Hartmann-Schröder, 1971: 207, fig. 67a–f. Eunereis longissima.—Sardá, 1983: 18, fig. 3a–f. Nereis longissima.—Chambers and Garwood, 1992: 35, figs. 5,11,27,34,44,56,67.
Material Examined.—One spec., sta. DI 21; one spec., sta. CM 27; one spec., sta. DI 26. Additional Material of Fauna Ibérica II.—One spec. (86 DL), Apr. 1991, west of Ría de Muros, 24°41.74´N, 09°10.06´W, depth 84 m (beginning), 42°41.60´N, 09°09.76´W, depth 81 m (end), sandy-mud with shells. One spec. (152 A), Apr. 1991, north of San Sebastián, 43°22.59´N, 02°03.70´W, depth 107 m (beginning), 43°22.16´N, 02°05.76´W, depth 104 m (end), black mud. Description.—Three incomplete specimens without posterior end. Largest specimen with 60 setigers, 18 mm long and 0.7 mm wide without parapodia and setae. Preserved specimens yellowish or brown in color, no pigmentation patterns. Prostomium subpyriform, frontal antennae slender, slightly larger than palps. Palps broad, with globular palpostyles. Two pairs of small eyes, arranged in square. Peristomium slightly longer than first setiger. Tentacular cirri slender, longest one extending back to setiger 11 (Fig. 6A). Pharynx with paragnaths only on oral ring, few in number, 1 in area VI; pair of brown serrated and curved jaws, with 13 teeth. Setigers 1–2 with uniramous parapodia, dorsal and ventral cirri shorter than lobes. Superior and inferior lobes triangular, setigerous lobe conical (Fig. 6B). Parapodia bi- ramous from setiger 3, dorsal cirri slender, shorter than lobes in anterior setigers, longer in posterior setigers. Upper and lower notopodial lobes triangular and subequal in size. Neuropodial acicular lobes conical, without prominent accesory lobes, short than notopodial and inferior neuropodial lobes. Neuropodial inferior lobes subequal in size and shape to notopodial lobes (Fig. 6C–E). Ventral cirri short and subulate. Notosetae few in number (2–3), in anterior setigers homogomph spinigers. In mid- body (setiger 15) and posterior region notopodia with 1–2 stout homogomph falcigers, with short, elongate-oval blades, smooth or serrated (3–4 teeth) (Fig. 6L, M). Neurosetae of upper bundle homogomph spinigers (1–5) and heterogomph falcigers (2–3) (Fig. 6F,H,K); lower bundle heterogomph spinigers (Fig. 6I) and falcigers, with gradation in length of blades of spinigers (2–3) to falcigers (2–3) (Fig. 6G, J). Heterogomph falcigers from upper and lower bundles similar in size, with long and slender, unidentate blades . Remarks.—The genus Eunereis Malmgren, 1867, based on the type-species Nereis longissima, Johnston, 1840, is characterized by having paragnaths only on the oral ring; these are occasionally difficult to observe and can be easily lost (Chambers and Garwood, 1992). The presence of paragnaths only on the oral ring distinguishes this genus from related genera, for example, Nereis, Neanthes and Ceratonereis. Another characteristic is the presence of homogomph falcigers among the notosetae from the middle of the body, NÚÑEZ ET AL.: NEREIDIDAE FROM CAPBRETON CANYON, BAY OF BISCAY 35
Figure 6. Eunereis longissima. A, anterior end, dorsal view, B. uniramous parapodium, setiger 2, C. biramous parapodium, setiger 6, D. parapodium, setiger 28, E. parapodium, setiger 47, F. falcigerous neuroseta, upper bundle, from setiger 6, G. falcigerous neuroseta, lower bundle, from setiger 6, H. falcigerous neuroseta, upper bundle, from setiger 28, I. spinigerous neuroseta, lower bundle, from setiger 28, J. falcigerous neuroseta, lower bundle, from setiger 28, K. falcigerous neuroseta, juvenile specimen, L. homogomph falcigerous notosetae from setigers 47 and 48, M. homogomph falcigerous notoseta, juvenile specimen. Scale: A = 0.6 mm; B–E = 206 µm; F–M = 29 µm. 36 BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000 which can replace the homogomph spinigers (Fauchald, 1972). The difficulty in observ- ing the paragnaths (sometimes yellowish), which can also become loosened, has led to numerous mis-identifications of species of other genera such as Rullierinereis Pettibone, 1971, a genus that is characterized for having a pharynx without any paragnaths and papillae, with the rest of the characters being similar to Eunereis. These specimens were small in size, but the adults can be large (315 mm length). However, variations in the size of the parapodial lobes and the blade length of homogomph falcigers are small. In our specimens we have observed very few conical paragnaths well sclerotised in areas VI, VII and VIII of the oral ring; these paragnaths were lost easily when manipulated. Another characteristic that can lead into incorrect determinations is that some specimens have eyes lacking pigmentation. Distribution.—Northeast Atlantic: from Norway to Bergen, 60°N, the Faroes and west coast of Europe to Mediterranean (10–2000 m) (Kirkegaard, 1998). Black Sea.
ACKNOWLEDGMENTS
We are very grateful to the crew of the RV CÔTE D’AQUITAINE for their helpful assistance at sea and the CIRMAT (INSU—CNRS) for logistical support and loan of an epibenthic dredge; to J. Corbera for providing the final versions of the species drawings; to T. Miura for the loan of speci- mens of Nicon sinica. This study was supported in part by the Dirección General de Enseñanza Superior de la Secretaría de Estado de Universidades e Investigación del Ministerio de Educación y Cultura, project research “Fauna Ibérica IV” (PB95-0235).
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ADDRESSES: (J.N.) Departamento de Biología Animal (Zoología), Facultad de Biología, Universidad de La Laguna, 38206 La Laguna, Tenerife, Canary Islands, Spain. E-mail: