Lumbrineridae ( Polychaeta ) from the Capbreton Canyon ( Bay of Biscay , NE Atlantic ) with the description of two new species

The Capbreton Canyon is a ’gouf’ type submarine valley which divides the continental shelf of the Bay of Biscay into two zones: the wide Aquitanian continental shelf, to the north, and the narrow Cantabrian shelf, to the south. Between 1987 and 1990, the CAPBRETON oceanographic cruises were undertaken, with one of the objectives of this programme being the characterisation of the faunal communities of the two levels defined by Le Danois (1948). Some of the results obtained on macrobenthic polychaete communities have been presented in Rallo (1988), Rallo et al. (1993a, b), García Arberas and Rallo (1994), San Martín et al. (1996), Aguirrezabalaga et al. (1999, 2001, 2002), Aguirrezabalaga and Ceberio (2003, 2005a, b, 2006) and Núnez et al. (2000). The objective of this study is to contribute to the knowledge of the polychaete communities of the SCIENTIA MARINA 70S3 December 2006, 17-25, Barcelona (Spain) ISSN: 0214-8358

SCIENTIFIC ADVANCES IN POLYCHAETE RESEARCH R. Sardá, G. San Martín, E. López, D. Martin and D. George (eds.)slope of the Capbreton Canyon, and especially of the family Lumbrineridae.
Commonly, lumbrinerids have been regarded as polychaetes with simple body shape and reduced external morphological characters.The species described were placed in a few genera and some genera were regarded as synonyms (Carrera-Parra, 2001).At present, the taxonomic delineation of the family is changing due to a different taxonomic approach based mainly on the use of maxillary characters, and some genera have been emended (Carrera-Parra, 2004).This work includes the description of eight species belonging to six genera, from which two are described as new species belonging to the genera Augeneria and Lumbricalus.

MATERIALS AND METHODS
The specimens were collected at 18 stations situated between 480 and 1113 m depth (Fig. 1 and Table 1).Samples were taken with three different types of gear: a Sanders-Hessler epibenthic dredge (DI) with a mesh size of 0.5 mm, a Flusha (KF) box corer (samples were sieved through a 0.5-mm-mesh screen), and a Marinovitch trawl (CM).Specimens were preserved in a 10% formaldehyde-seawater solution.
The descriptions are based mainly on the best specimen.An anterodorsal dissection was made to extract the maxillary apparatus; it was mounted dorsally and ventrally on a slide in order to examine details of both the maxillae and the mandible under a compound microscope.A true tooth is herein restricted to those structures provided with a pulp cavity.The maxillae are abbreviated as M and followed by a roman numeral indicating the position in the maxillary apparatus from dorsal to ventral side.The measurements were standardized for length through chaetiger 10 (L10), and for width at chaetiger 10 excluding parapodia (W10).Illustrations were made using camera lucida and some digital photographs were taken through a light microscope.Some type material was borrowed from the collections of the Muséum National

SYSTEMATICS
Family LUMBRINERIDAE Schmarda, 1861, emended Orensanz, 1990Genus Abyssoninoe Orensanz, 1990 Abyssoninoe scopa (Fauchald, 1974) Lumbrineris scopa Fauchald, 1974: 26-27, Fig. 5. Description.Specimen incomplete with 174 chaetigers, L10= 3.0 mm, W10= 1.12 mm.Prostomium conical, slightly longer than wide, with a pair of prominent nuchal organs, with short palps ventrally.Peristomium shorter than prostomium, with two rings; anterior ring slightly longer than the second one; separation between rings distinct dorsally and laterally; ventrally the first ring incom-plete, second one projected forward as a muscular lip.All parapodia well developed, first seven smaller than following ones.Prechaetal lobe inconspicuous along the body.Postchaetal lobe well developed from the first parapodium, conical, becoming more developed, digitiform, in posterior parapodia.Short rounded dorsal cirri in all parapodia, with several notoaciculae.Limbated simple multidentate hooded hooks in chaetigers 1-20; simple multidentate hooded hooks with long hood in chaetigers 20-32, with up to 8 teeth of similar size; from chaetiger 33 with short hood, with up to 8 teeth, proximal tooth bigger.Aciculae yellow, aristate, up to four in anterior parapodia, and one in posterior parapodia.Maxillary apparatus with four pairs of maxillae; maxillary carriers slightly shorter than MI, anterior end constricted; MI forceps-like, with bridles well developed; MII as long as MI, with four teeth in left plate and five in the right one.MIII arcuate, unidentate; MIV unidentate, like a broad rectangular plate with a projection from middle of its inferior border.Mandible fused for up to 3/4 of its length.

Material examined. Atlantic Ocean, Bay of Biscay, Capbreton Canyon (coordinates in
Remarks.Additional material L10 = 1.30-3.30mm, W10 = 0.25-1.15mm.The ending of limbated simple multidentate hooded hooks varies from chaetigers 9 to 21, and the beginning of simple mul-  tidentate hooded hooks with long hood in chaetiger and simple multidentate with short hood vary from chaetigers 10 to 22 and 23 to 41 respectively; all are size dependent. Discussion.Parapar et al. (1994) considered Abyssoninoe scopa (Fauchald, 1974), among other species, as a synonym of A. hibernica (McIntosh, 1903).However, herein, we are proposing that there are enough important differences, mainly in the development of the postchaetal lobes at the posterior end, to consider both as valid species.A review and comparison of complete specimens of all Northeast Atlantic species is necessary to clarify the taxonomy of these groups of species.
Genus Augeneria Monro, 1930, emended Orensanz, 1973 Augeneria algida (Wirén, 1901) Augeneria algida Winsnes, 1987:40, Figs Description.Specimen incomplete with 27 chaetigers, L10 = 1.7 mm, W10 = 0.5 mm.Prostomium conical, slightly longer than wide, with a pair of nuchal organs, with three small antennae in a single row; with short palps ventrally.Peristomium shorter than prostomium, about 1/2 its length; with two rings, anterior ring slightly longer than the second one; separation between rings distinct dorsally and laterally; ventrally first ring incomplete, second one projected forward as a muscular lip.All parapodia well developed, first five smaller than following ones, slightly visible in dorsal view.Prechaetal lobe inconspicuous along the body.Postchaetal lobe well developed from first parapodium, conical, slightly longer than prechaetal lobe, more developed in parapodia 6-13.Short rounded dorsal cirri in all parapodia.Composite multidentate hooded hooks in chaetigers 1-12, with long blade, with up to 7 teeth of similar size; simple multidentate hooded hooks from chaetiger 13, with up to 7 teeth, proximal tooth bigger, with short hood; dorsal limbates in all chaetigers of the specimen, ventral limbates in chaetigers 1-14.
Aciculae yellow, aristate, up to three in anterior parapodia, and one in posterior one.Maxillary apparatus with four pairs of maxillae; maxillary carriers slightly shorter than MI; MI forceps-like with bridles well developed; MII as long as MI, with three stout teeth.MIII arcuate, edentate; MIV edentate, with whitish central area.Mandible with both anterior and posterior end divergent.
Remarks.Additional material L10 = 1.20-1.52mm, W10 = 0.25-0.39mm.The ending of composite multidentate hooded hooks and the beginning of the simple multidentate hooded hooks varies from chaetiger 8 to 12 and 9 to 13 respectively; both are size dependent.
Description.Holotype incomplete with 29 chaetigers, L10 = 8.2 mm, W10 = 4.5 mm.Prostomium conical, as long as wide, with a pair of prominent nuchal organs, with eight small antennae in two rows (Fig. 2A); with well developed palps ventrally.Peristomium as long as the prostomium; with two rings, anterior ring twice as long as the second one; separation between rings distinct dorsally and laterally; ventrally first ring incomplete, second one projected forward as a muscular lip, transversally separated.All parapodia well developed, first five smaller than following ones.Prechaetal lobe in parapodia 1-7 inconspicuous, from parapodium 8 with a small digitiform projection in the distal end increasing slightly.in parapodia 13-19 similar in size to the postchaetal lobe, from parapodium 20 slightly longer than postchaetal lobe (Figs.2B-C).Postchaetal lobe well developed in all parapodia, in parapodia 1-4 conical, from parapodium 5 with a digitifom projection increasing slightly, longer than prechaetal lobe, from parapodium 20 shorter than prechaetal one.Short rounded dorsal cirri in all parapodia, with several notoaciculae.Composite multidentate hooded hooks in chaetigers 1-19, with short blade, with up to 7 teeth of similar size (Fig. 2D); simple multidentate hooded hooks from chaetiger 20, with short hood, with up to 8 teeth of similar size (Fig. 2E); dorsal and ventral limbates in all chaetigers of the specimen.Aciculae dark, aristate, up to five in anterior parapodia, and three in posterior ones.Maxillary apparatus with four pairs of maxillae (Fig. 2F); maxillary carriers shorter than MI; MI forceps-like with bridles well developed; MII as long as MI, with three stout teeth, the basal one poorly developed.MIII arcuate, edentate; MIV edentate, with whitish central area.Mandible with both anterior and posterior end divergent.
Etymology.This species is named in honour of Enrique Rioja in recognition of his notable studies in invertebrate zoology, especially his studies on polychaetes from Spain and Mexico.(Fig. 3)  Description.Holotype incomplete, with 150 chaetigers, L10 = 7.0 mm, W10 = 4.0 mm.Prostomium conical, rounded, as long as wide, with a pair of nuchal organs, with well developed palps ventrally.Peristomium half as long as prostomium, with two rings; anterior ring twice as long as the second one; separation between rings distinct dorsally and laterally, ventrally first ring incomplete and second one projected forward as a muscular lip.All parapodia well developed, first 10 smaller than following ones.Prechaetal lobe inconspicuous along the body.Postchaetal lobe well developed in all parapodia; in parapodia 1-4 conical, from parapodium 5 to 30 conical, wide basally; from parapodium 41 digitiform; always longer than the prechaetal lobe (Figs.3A-C).Parapodia and lobes more developed from parapodium 10 to 23.With conspicuous short dorsal cirri in all parapodia, with several notoaciculae.Composite spinigers in chaetigers 1-28 (Fig. 3D), with up to two or three per parapodium; composite multidentate hooded hooks in chaetigers 1-30, with long blade, with up to 10 teeth of similar size (Fig. 3E); simple multidentate hooded hooks from chaetiger 31, with up to 5 teeth, proximal tooth bigger, with short hood (Fig. 3F); dorsal limbates in chaetigers 1-107, ventral limbates in chaetigers 1-35.All chaetae reddish.Aciculae black, aristate, up to five in anterior parapodia, and two in posterior parapodia.Maxillary apparatus with five pairs of maxillae (Fig. 3G).Maxillary carriers shorter than MI, anterior end constricted.MI forceps-like, with well developed bridles; MII shorter than MI, with wide well developed connecting plates, with 4 teeth.MIII unidentate, with prominent tooth followed by an expanded base; MIV unidentate; MV free, lateral to MIV and MIII.Mandible fused for up to 3/4 of its length.Etymology.This species is named in honour of Antonio Campoy in recognition of his studies on polychaetes from the Iberian Peninsula.

Discussion
Discussion.L. campoyi n. sp.resembles L. harrisae Carrera-Parra, 2004from California, L. composita (Hartmann-Schröder, 1965) from Chile, and L. aotearoae (Knox and Green, 1972) from New Zealand, by the presence of MIII unidentate.It differs from L. harrisae and L. aotearoae by the colour of the aciculae, black in L. campoyi n. sp. and yellow in the other two species.L. campoyi n. sp.differs from L. composita because it lacks elongated postchaetal lobes from medium parapodia.Furthermore, L. campoyi n. sp. is the only species of this group having few composite spinigers per parapodium (2-3).
Distribution.Restricted to type locality.
Description.Specimen incomplete, with 34 chaetigers, L10 = 2.5 mm, W10 = 0.4 mm.Prostomium three times longer than wide, with a small pair of nuchal organs, with short palps ventrally.Peristomium shorter than prostomium, with two rings of similar size; separation between rings distinct dorsally and laterally; ventrally first ring incomplete and second one projected forward as a muscular lip.First 5 parapodia smaller than following ones, slightly visible in dorsal view.In parapodia 1-6 both prechaetal and postchaetal lobes rounded, short, of similar size.From parapodium 7 postchaetal lobe more developed, digitiform throughout the body.Parapodia and lobes more developed from parapodium 9 to 16.Short dorsal cirri in all parapodia.Simple bidentate hooded hooks from chaetiger 5; limbates in all chaetigers of the specimen.Aciculae yellow, aristate, with up to three in anterior parapodia and two in posterior parapodia.Maxillary apparatus with four pairs of maxil-lae; carriers as long as MI, slightly constricted in the middle region.MI forceps-like with internal accessory teeth, with well developed bridles; MII as long as MI, with 3 teeth; MIII arcuate, unidentate; MIV unidentate with attachment lamella slightly developed.Mandible fused, reddish.
Distribution.Northeast Atlantic (Spain and France).

Genus Lumbrineris de Blainville, 1828
Lumbrineris aniara Fauchald, 1974 Lumbrineris aniara Fauchald, 1974: 24-25 Description.Specimen incomplete with 88 chaetigers, L10 = 2.7 mm, W10 = 0.8mm.Prostomium conical, as long as wide, with a pair of nuchal organs, with well developed palps ventrally.Peristomium slightly shorter than prostomium, with two rings of similar size; separation between rings distinct dorsally and laterally; ventrally the first ring incomplete and the second one projected forward as a muscular lip.All parapodia well developed, first four smaller than following ones.Prechaetal lobe in parapodia 1-6 inconspicuous, from parapodium 7 short, rounded throughout the body.Postchaetal lobe well developed from first parapodium; in parapodia 1-18 auricular, from parapodium 19 digitiform, always longer than prechaetal lobe.Short dorsal cirri in all parapodia.Composite multidentate hooded hooks in chaetigers 1-13, with short blade, with up to 6 teeth all of similar size; simple multidentate hooded hooks from chaetiger 14, with up to 8 teeth, proximal tooth slightly bigger, with short hood; dorsal limbates in chaetigers 1-48, ventral limbates in chaetigers 1-15.Aciculae yellow, aristate, up to three in anterior parapodia, and one in posterior parapodia.Maxillary apparatus with five pairs of maxillae; maxillary carriers shorter than MI, almost constricted in the middle region.MI forceps-like with well developed bridles.MII as long as MI, with 4 teeth.MIII unidentate, with short tooth; MIV with prominent tooth.MV free, lateral to MIV and MIII.Mandible divided for about half its length.
Remarks.Additional material L10 = 1.40-2.70mm, W10 = 0.27-0.85mm The ending of composite multidentate hooded hooks and the beginning of the simple multidentate hooded hooks varies from chaetiger 6 to 15 and 7 to 16 respectively; both are size dependent.
Distribution.Northeast Atlantic.Kinberg, 1865 Lumbriconereis futilis Kinberg, 1865: Description.Specimen incomplete with 84 chaetigers, L10= 3.0 mm, W10= 1.0 mm.Prostomium conical, as long as wide, with a pair of nuchal organs, with well developed palps ventrally.Peristomium shorter than prostomium, about 1/2 its length; with two rings; anterior ring twice as long as second one; separation between rings distinct dorsally and laterally; ventrally the first ring incomplete and the second one projected forward as a muscular lip.All parapodia well developed, first four smaller than following ones.Prechaetal lobe in parapodia 1-15 inconspicuous, from parapodium 16 slightly developed, rounded.Postchaetal lobe well developed from the first parapodium; in parapodia 1-4 digitiform, in parapodia 5-16 digitiform, wide basally; from parapodium 17 digitiform, always longer than prechaetal lobe.Parapodia and lobes better developed in parapodia 7-14.Short dorsal cirri in all parapodia.Composite multidentate hooded hooks in chaetigers 1-17, with long blade, with up to 7 teeth, all of similar size; simple multidentate hooded hooks in chaetigers 18-25, with long hood, with up to 7 teeth, proximal tooth bigger, from parapodium 26, with short hood and with up to 7 teeth, proximal tooth bigger.Aciculae black, aristate, up to three in anterior parapodia, and one in posterior parapodia.Maxillary apparatus with five pairs of maxillae; maxillary carriers slightly shorter than MI, anterior end constricted.MI forceps-like with well developed bridles; MII as long as MI, with 4 teeth.MIII unidentate, with a prominent tooth followed by a knob; MIV with pointed tooth; MV free, prominent, lateral to MIV and MIII.Mandible divided for about half its length.

Lumbrineris futilis
Discussion.Hartman (1949) considered this species a synonym of Lumbrineris latreilli, but there are sufficient differences between them to consider both species as valid; the main differences including the number of teeth in MIII and the colour of the aciculae.
Description.Specimen incomplete with 77 chaetigers, L10= 4.0 mm, W10= 1.1 mm.Prostomium conical, twice as long as wide, with a pair of divergent dorsal longitudinal black bands, with a pair of nuchal organs, with well developed palps ventrally.Peristomium shorter than prostomium, about 1/2 its length; with two rings; anterior ring slightly longer than the second one; separation between rings distinct dorsally and laterally; ventrally first ring incomplete and second one projected forward as a muscular lip.All parapodia well developed, first five smaller than following ones.Prechaetal lobe inconspicuous in all parapodia; postchaetal lobes well developed from first parapodium; in parapodia 1-36 digitiform, more developed from parapodium 6 to 36, shorter from parapodium 36.Small dorsal cirri in all parapodia, with several notoaciculae.Branchiae from parapodium 3 to 36, with up to five branchial filaments.Simple multidentate hooded hooks in chaetigers 7-36, with long hood, with up to 8 teeth, all of similar size; from parapodium 37, with short hood and with up to 7 teeth, proximal tooth bigger; limbates in all chaetigers of the specimen.Aciculae reddish, aristate, with up to three in anterior parapodia and two in posterior one.Maxillary apparatus with five pairs of maxillae; maxillary carriers as long as MI, constricted in anterior end; MI forceps-like, with bridles well developed; MII as long as MI; with 7 teeth; MIII with 4 teeth, distal tooth bigger; MIV with 16 teeth, distal tooth bigger; MV free, lateral to MIV and MIII.Mandible fused for up to 3/4 of its length.

Distribution.
Northeast Atlantic and Mediterranean.
. Augeneria riojai n. sp.resembles A. polytentaculata Imajima and Higuchi, 1975 from Japan by having several small antennae and by the shape and development of both the prechaetal and postchaetal lobes; it differs mainly because Augeneria riojai n. sp. has dark aciculae and A. polytentaculata has yellow aciculae.Type locality.Capbreton Canyon, Bay of Biscay.Distribution.Restricted to type locality.Genus Lumbricalus Frame, 1992, emended Carrera-Parra, 2004 Lumbricalus campoyi n. sp.