Published in collaboration with the University of Bergen and the Institute of Marine Research, Norway

Two new genera of the pelagic family Yndolaciidae (Polychaeta) from the Arctic Ocean with an addition to the description of Yndolacia lopadorrhynchoides Støp-Bowitz

Galina N. Buzhinskaja SARSIA Buzhinskaja GN. 2004. Two new genera of the pelagic family Yndolaciidae (Polychaeta) from the Arctic Ocean with an addition to the description of Yndolacia lopadorrhynchoides Støp-Bowitz. Sarsia 89:338–345.

Until recently, the pelagic family Yndolaciidae was considered to include a single genus and species Yndolacia lopadorrhynchoides Støp-Bowitz, 1987 from the Gulf of Guinea. Yndolaciella polarsterni gen. et sp. nov. and Paryndolacia tomopteroides gen. et sp. nov. are described from the deep sea of the Arctic Ocean. Additions and refining of the diagnoses of the genus Yndolacia and the family are given. The absence of cephalic tentacles, a weakly developed brain, simple biramous parapodia with equal rami, similar cirri without cirrophores, a superficial position of the nervous system, the presence of two nerve cords and the lack of ganglionic swellings are assumed to constitute ancestral characters of the family. Some resemblance of the family to Tomopteridae is noted. It is supposed that fossil Eotomopteris aldridgei Briggs & Clarkson, 1987 belongs to Yndolaciidae.

Galina N. Buzhinskaja, Zoological Institute, Russian Academy of Sciences, St. Petersburg 199034, Russia. E-mail: [email protected]

Keywords: Polychaeta; Yndolaciidae; new genera; Arctic Ocean.

INTRODUCTION MATERIAL AND METHODS

The pelagic polychaete family Yndolaciidae was The specimens of Yndolacia lopadorrhynchoides were erected to comprise Yndolacia lopadorrhynchoides collected in 1960–1961, during the 13th and 14th Støp-Bowitz, 1987 from the Gulf of Guinea. The cruises of R/V Ombango, from the Gulf of Guinea. authority of the family wrote: “The reduced prosto- Open net Grand Smidt (vertical hauls, net opening mium and the enormous tentacular cirri provided 9m2, mesh size 1 mm) was used to collect the plankton with chaetae set this apart from any other samples. All of the species were found in known form … Yndolacia should be placed in a bathyplankton from depth 725–880 m in intermediate separate family, Yndolaciidae, provisionally without Antarctic water (Støp-Bowitz 1992). The seawater obvious relationships to any other forms” (Støp- temperature at depths of 800–880 m in June 1960 was Bowitz 1987). Fauchald & Rouse (1997) considered 4.7–4.9 °C, salinity 34.5–34.6%. A macerated speci- Yndolaciidae among “poorly known and poorly men of Yndolacia lopadorrhynchoides was used to understood polychaete families”. Muir (in Chambers draw the external view of the nervous system. The & Muir 1997) placed Yndolaciidae in the new order material is deposited at the Museum National d’Histoire Yndolaciida. Naturelle (Paris, France). Downloaded by [TOBB Ekonomi Ve Teknoloji] at 13:30 21 December 2014 Brief information about the first finding of animals Yndolaciella polarsterni gen. et sp. nov. was brought, belonging to the family in the Arctic has been given among other invertebrates, on board R/V Polarstern in without description of the specimens (Buzhinskaja in September 1993, from the Laptev Sea, by an open Sirenko & al. 1996; Buzhinskaja 2001a, b). In this benthopelagic sampler attached to an Agassiz trawl. paper, two new genera and species are described, based The frame opening of the sampler was 25Â95 cm, the on that material. Examination of two specimens mesh size of the net was 0.4 mm. Details concerning the of Yndolacia lopadorrhynchoides from the Gulf of gear were given by Sirenko & al. (1996). Catching was Guinea, kindly provided by Dr Dauvin, allowed the started near the bottom at a depth of about 3040 m. diagnosis of Yndolacia to be defined more precisely. A seawater temperature of À0.755 °C and salinity of Some additions to the description of the family 34.88% were measured at a depth of 3065 m. The and comparisons with some other are sample was preserved in 4% formaldehyde and later given. transferred to 70% alcohol.

DOI 10.1080/00364820410002604 # 2004 Taylor & Francis Buzhinskaja – Two new genera of Yndolaciidae (Polychaeta) from the Arctic Ocean 339

Paryndolacia tomopteroides gen. et sp. nov. was Remarks obtained from a depth of 3290 m during the drift of the Russian drifting station North Pole 22 in the Canada Støp-Bowitz (1987: 128, figs 1, 2) stated that the Basin in 1977. A small Sigsbee trawl with a frame prostomium of Yndolacia is highly reduced and that opening of 15Â80 cm was employed for sampling. A nuchal organs are located on the peristomium. More- temperature of À0.4 °C, salinity of 35% and a high over, the prostomium is lacking in his drawings. But concentration of oxygen were characteristic of the deep two specimens re-examined have a distinct prosto- water of the Canada Basin (Levenstein 1981). The mium, with nuchal organs on its posterior part. As species was found together with polychaetes of the distinguished from Støp-Bowitz’s figures, where tenta- subfamily Macellicephalinae. cular cirri of the first segment are shorter than those Both new species are deposited at the Zoological of the second, the tentacular cirri of the specimens Institute of the Russian Academy of Sciences, St. examined are subequal. Emergent terminal chaetae on Petersburg (ZIRAS). The drawings were made by the tentacular cirri mentioned by Støp-Bowitz are means of a Karl Zeiss light microscope with a mirror absent. Besides, all cirri of the animals are broad at attachment. the base and have no distinct cirrophores. Yndolacia lopadorrhynchoides Støp-Bowitz, 1987 (Fig. 1) Yndolacia lopadorrhynchoides Støp-Bowitz, 1987: Order Phyllodocida 128–130, figs 1–4; 1992: 34–36, fig. 10 Family Yndolaciidae Støp-Bowitz, 1987 Material examined Diagnosis One specimen, female with eggs. Gulf of Guinea Body short, flattened, semitransparent. Prostomium (1°55'S8°30'E), R/V Ombango, cruise 13, Stn 309, distinct, broad, with one pair of ciliated nuchal organs. 17 June 1960, about 800 m. One poorly preserved Palps, antennae and eyes absent. First segment enclos- specimen. Gulf of Guinea (9°08'S7°00'E), R/V ing ventral mouth, reduced dorsally and partly fused Ombango, cruise 14, Stn 321, 28 February 1961, about with prostomium. Each of two anterior segments with a 725 m. pair of voluminous tentacular cirri supported by inner chaetae. Subsequent segments with biramous parapodia Additions to description having long stem and equal rami, each supported by single acicula. Dorsal and ventral cirri large, equal, Length of female about 10 mm, width between para- elongated or oval. Chaetae thin, compound. Branchia podia about 2.3 mm, width including parapodia about absent. Proboscis short, unarmed. Pygidium with or 3.9 mm. Body flattened, semitransparent, about 20 without a pair of short anal cirri. Two closely arranged segments. Prostomium short and wide without eyes, nerve cords without visible ganglionic swellings. palps or antennae, with a pair of multiramified ciliated Pelagic forms. nuchal organs (Fig. 1A). Proboscis retracted (not Type genus: Yndolacia Støp-Bowitz, 1987 examined). First segment enclosing ventral mouth reduced and not visible at dorsal side, completely Yndolacia Støp-Bowitz, 1987 coalesced with prostomium and second segment. Two pairs of subequal voluminous tentacular cirri on two Downloaded by [TOBB Ekonomi Ve Teknoloji] at 13:30 21 December 2014 Diagnosis anterior segments; each cirrus broad at the base, tapering distally and supported by one to two fine short About 20 segments. Prostomium short, wide. Ramified chaetae and two larger chaetae (aciculae?). Tentacular nuchal organs, situated dorsally on posterior part of cirri about as long as width of body. Biramous prostomium. First segment reduced, completely fused parapodia with elongated stem ending in two pointed with prostomium and second segment. Tentacular cirri branches (Fig. 1B, C). Noto- and neuropodium sub- long subequal, with broad base; supported by a few equal, each supported by long acicula. Long subequal inner chaetae. Parapodia with numerous chaetae. dorsal and ventral cirri bottle-shaped, broadly based and Acicular lobes elongated, pointed. Podial cirri bottle- tapering to a point. Podial cirri about as long as length shaped, pointed. Pygidium unknown. of podial stem. Both rami having many compound thin Type species: Y. lopadorrhynchoides Støp-Bowitz, chaetae with long slender blades and heterogomph 1987. articulations (Støp-Bowitz 1987: fig. 4). 340 Sarsia 89:338-345 – 2004

Fig. 1. Yndolacia lopadorrhynchoides Støp-Bowitz. A. Anterior end, dorsal view; the dotted line shows the posterior border of the first segment. B. Parapodium (Stn 309). C. Parapodium (Stn 321). D. Circumpharyngeal connectives and nerve cords, ventral view. Scale bars = 1 mm (D); 0.5 mm (A, B); 0.2 mm (C). Downloaded by [TOBB Ekonomi Ve Teknoloji] at 13:30 21 December 2014

Ventral nerve cords close to each other; ganglia not Diagnosis visible (Fig. 1D). Pygidium unknown. Female contains oval eggs (up to 40Â70 mm) in coelom. Body short, nine segments. Prostomium broad with a pair of simple ciliated nuchal organs at posterior margin. First segment fused with prostomium, partly visible dorsally. Tentacular cirri supported by thin inner Remark chaetae. Noto- and neuropodium each with two to three compound chaetae; acicular lobes short. Dorsal and The autapomorphic characters of Yndolacia: fusion of ventral cirri long, not morphologically separated from first and second segments and elaborate nuchal organs. podial rami. Pygidium with two short anal cirri. Anus Yndolaciella gen. nov. subdorsal. Buzhinskaja – Two new genera of Yndolaciidae (Polychaeta) from the Arctic Ocean 341

Type species: Y. polarsterni sp. nov. in the different shape and length of podial cirri. As distinguished from Yndolacia, two anterior segments of Yndolaciella polarsterni sp. nov. (Fig. 2) this species are distinctly separated from each other and Yndolaciidae gen. et sp. nov. Buzhinskaja, in Sirenko podial cirri are not demarcated morphologically from & al. 1996: 348; Yndolaciella polarsterni Buzhinskaja podial rami. Besides, Yndolaciella has the simple nuchal 2001b: 53 (nomen nudum). organs and a small number of chaetae in parapodia. The autapomorphic character of Yndolaciella is a Type material reduced number of segments. Holotype: ZIRAS No. 1/50433. Paryndolacia gen. nov.

Type locality Diagnosis ° ' ° ' Laptev Sea (79 11.3 N 119 56.4 E), 3042–0 m, RV About 20 segments. Prostomium unknown. Noto- and Polarstern, Stn 54, 13 September 1993. Coll. B. I. neuropodium with two to three compound chaetae; Sirenko. acicular lobes short leaf-like. Podial cirri oval, short, voluminous, morphologically not separated from podial Etymology rami. Pygidium without cirri. Anus ventral. The specific name polarsterni refers to the research vessel Polarstern, which collected the specimen. Remark

Description Even though the anterior end of the genus is unknown, the general plan of the parapodial structure and Length 2.1 mm, width between parapodia 0.3 mm. similar chaetae allow it to be placed in the family Body flattened, semitransparent; consisting of nine Yndolaciidae. segments (Fig. 2A). Prostomium wide, short with a pair of oval ciliated nuchal organs at posterior margin Paryndolacia tomopteroides sp. nov. (Fig. 3). (Fig. 2B). Retracted proboscis short, without jaws. First P. tomopteroides Buzhinskaja 2001b: 53 (nomen segment fused with prostomium, reduced dorsally, but nudum). its posterior margin visible dorsally; separated from second segment (Fig. 2B). First and second segments Type material each with a pair of stout tentacular cirri of nearly equal length, which is 7 times longer than width of body. Holotype: ZIRAS No. 1/50432 (fragment). Broadly based tentacular cirri more or less abruptly narrowing into thread-like appendages, supported by Type locality two to three thin simple chaetae (Fig. 2A, B). Sub- Arctic Ocean, Canada Basin (79°26.2'N 127°21.2'W), sequent segments with biramous parapodia. Podial stem 3290 m, expedition North Pole 22, Stn 34, 1971. elongated. Noto- and neuropodium equal, each having short leaf-shaped pointed acicular lobe and elongated Etymology cirrus. Aciculae comparatively long: up to middle of podial stem. Dorsal and ventral cirri subequal, markedly The specific name tomopteroides is given due to a longer than podial stem, broadly based and gradually superficial likeness with Tomopteris.

Downloaded by [TOBB Ekonomi Ve Teknoloji] at 13:30 21 December 2014 tapering to end; morphologically not separated from podial rami (Fig. 2C). Chaetae thin, compound with Description slender smooth blades and heterogomph articulations; One fragment consisting of 17 chaetigerous segments one to three in each ramus (Fig. 2D). and pygidium (Fig. 3A). Body flattened. Length Pygidium with two short, basally expanded, anal cirri 6.8 mm, greatest width including parapodia about (Fig. 2E). Anus subdorsal. 3 mm, width between parapodia about 1.3 mm. Colour Nerve cords close to each other; ganglia not visible yellowish, semitransparent. (Fig. 2A). Parapodia with long stem. Noto- and neuropodia Remarks equal, each supported by light acicula curved distally (Fig. 3B). Aciculae relatively short, up to beginning of Yndolaciella differs from other species of Yndolaciidae podial stem. Acicular lobes short, leaf-like, pointed. in the short body with a small number of segments and Dorsal and ventral cirri oval, voluminous, not separated 342 Sarsia 89:338-345 – 2004 Downloaded by [TOBB Ekonomi Ve Teknoloji] at 13:30 21 December 2014 Fig. 2. Yndolaciella polarsterni gen. et sp. nov. A. Habitus, ventral view; nerve cords and partly circumpharyngeal connectives are shown in the middle. B. Anterior end, dorsal view. C. Parapodium. D. Chaeta. E. Posterior end, dorsal view. Scale bars = 0.5 mm (A); 0.2 mm (B, E); 0.1 mm (C).

from podial rami. All cirri about three times shorter Remarks than podial stem. Chaetae thin, compound with long slender blades and heterogomph articulations; one to The form of the parapodia with a very long stem and three in each ramus (Fig. 3C). Pygidium without cirri short oval cirri is the character that clearly differentiates (Fig. 3D). Anus ventral. Ventral nerve cords close to Paryndolacia tomopteroides from other yndolaciids. each other; ganglia not visible (Fig. 3A). The species is also characterized by curved and Buzhinskaja – Two new genera of Yndolaciidae (Polychaeta) from the Arctic Ocean 343

Fig. 3. Paryndolacia tomopteroides gen. et sp. nov. A. Habitus, ventral view, nerve cords are shown in the middle. B. Parapodium. C. Chaeta. D. Posterior end, ventral view. Scale bars = 1 mm (A); 0.45 mm (D); 0.1 mm (B).

relatively short aciculae. In addition, Paryndolacia no cephalic appendages. A loss of cephalic tentacles differs from Yndolaciella in the absence of anal cirri due to a planktonic mode of life is improbable, and, unlike Yndolacia, it has a small number of chaetae. suggesting it represents an ancestral state. Nuchal

Downloaded by [TOBB Ekonomi Ve Teknoloji] at 13:30 21 December 2014 Podial cirri of Paryndolacia are not demarcated from organs are posterior dorsal prostomial processes (sim- podial rami like those of Yndolaciella, but not identical ple or complicated). They are well developed and in form. reminiscent of those in Notophyllum (Phyllodocidae). The autopomorphic characters of Paryndolacia are As distinguished from Yndolaciidae, nuchal organs in short dorsal and ventral cirri and aciculae distally Notophyllum are fused to the cirrophores of the first pair curved. of tentacular cirri (Pleijel 1991). The first segment in Yndolaciidae is dorsally reduced and partly fused with the cephalic lobe. In Yndolacia,it DISCUSSION is also fused to the second segment, which completely The prostomium in the yndolaciids is a well-differ- covers the first segment and posterior part of the entiated simple lobe. Unlike other polychaetes of the prostomium (Fig. 1A). These two segments each carry order Phyllodocida, prostomia of the Yndolaciidae have a pair of enormous tentacular cirri supported by thin 344 Sarsia 89:338-345 – 2004

inner chaetae. Two pairs of tentacular cirri with thin two pairs of long tentacular cirri with inner chaetae and inner chaetae on two anterior segments also occur in parapodia supported by aciculae. larvae of tomopterids, for instance in Tomopteris The position of the nervous system in the body wall helgolandica (A˚ kesson 1962). However, during later is superficial. It consists of two closely arranged cords development, the enormously elongated chaetae without visible ganglionic swelling, although it is not replace the thin larval chaetae in the second pair of known exactly that ganglia are completely lacking in tentacular cirri in Tomopteris; the chaetae in the first the ventral cords without a histological study. The one or even the whole first pair of cirri may be lost. absence of segmental swellings also occurs in Tomop- Only in some species of Tomopteridae the first pair of teridae (A˚ kesson 1962) and among bottom polychaetes tentacular cirri is well developed in adults, but these in Polygordiidae, Protodrilidae, Saccocirridae (Fraipont cirri are always much shorter than the cirri of the second 1884; Kotikova 1973; Bubko 1981), (Per- pair. According to A˚ kesson (1962), thin inner chaetae in eyaslawzeva 1896; Bubko 1984), Oweniidae (Bubko & tentacular cirri of Tomopteris are transitory larval Minichev 1972), but it is not characteristic of the structures. The formation of these structures differs from majority of polychaetes. Among the above-mentioned the normal development of chaetae in polychaetes. families, Polygordiidae (Fraipont 1884) and Oweniidae In 5–6-day-old larvae of Tomopteris helgolandica, the (Bubko & Minichev 1972; Lagutenko 1986) possess cirri of the first segment are arranged at the same level an unpaired ventral nerve cord. In saccocirrids, nerve as the mouth. Later, these cirri move in an anterior cords are moved more widely than in yndolaciids, direction and have not only a pre-oral but also a pre- tomopterids and protodrilids. cerebral position (A˚ kesson 1962). In yndolaciids, the The lack of eyes and sensory organs such as palps first pair of tentacular cirri is situated on the ventral and antennae suggests that the brain in yndolaciids is side slightly behind the mouth. It must be noted that weakly developed. This is confirmed by insignificant the general appearance of Yndolaciella is reminiscent thickening of the anterior part of circumpharyngeal of 10–12 day larvae of Tomopteris helgolandica due to connectives (Fig. 1D). The external view of the long tentacular cirri, the reduction of the first segment yndolaciids brain is reminiscent of the simple brain and the shape of parapodia. of Oweniidae (Bubko & Minichev 1972) and larvae of Støp-Bowitz (1987) paid attention to some resem- some polychaetes. The brain of tomopterids is more blance of biramous parapodia in the monogeneric developed (Hanstro¨m 1928; A˚ kesson 1962). family Lacydoniidae and in Yndolaciidae. Any lobes Yndolaciidae is still a poorly known family and in (except acicular tubercles), papillae, branchiae are in my opinion to raise the family to the rank of a separate fact lacking on the parapodia in both families. In order is premature. I believe that the family should be contrast to the yndolaciids, the parapodia of Lacydonia retained in the order Phyllodocida. Yndolaciidae possess have a very short stem, small cirri and simple capillary some primitive peculiarities. The absence of cephalic notochaetae. Besides, biramous parapodia in nine tentacles and the weakly developed brain, the simple known species of Lacydonia are asymmetrical: noto- biramous parapodia with equal rami (in size and struc- podia are shorter. ture) and the similar cirri without cirrophores, the combi- Støp-Bowitz (1987) evidently considered the broad nation of such features as superficial position of the base of tentacular and podial cirri of Yndolacia as nervous system, the presence of two nerve cords and the cirrophores. Distinct cirrophores were not found in the lack of ganglionic swellings constitute ancestral char- material examined. Moreover, podial cirri of Yndola- acters of the family. The resemblance in morphology ciella and Paryndolacia are not demarcated morpholo- of yndolaciids to tomopterids may be a result of their Downloaded by [TOBB Ekonomi Ve Teknoloji] at 13:30 21 December 2014 gically from podial rami and look like continuations of relationship, with the Yndolaciidae being more primitive the latter. This peculiarity and the long podial stem than the Tomopteridae. New findings of yndolaciids define some external similarity of parapodia of these and more detailed morphological studies are required to two genera and that of the tomopterids. It especially clarify the degree of similarity of these two families. concerns Paryndolacia, which has relatively short cirri. However, parapodia of Tomopteridae have neither ACKNOWLEDGEMENTS aciculae nor chaetae, which apparently were lost in the process of evolution. In this connection it should be The work was performed with the financial support of the noted that the fossil Eotomopteris aldridgei described Russian Foundation for Basic Research project 01-04-49648 and Grant on Biodiversity No. 89. I am indebted to Dr D. by Briggs & Clarkson (1987) as a tomopterid from the Eibye-Jacobsen for critical comments and valuable discussion. Lower Carboniferous of Scotland, possibly belongs to I am also grateful to Dr A. Muir and an anonymous referee for Yndolaciidae. Like the yndolaciids, Eotomopteris has the constructive critical comments on the manuscript. Buzhinskaja – Two new genera of Yndolaciidae (Polychaeta) from the Arctic Ocean 345

REFERENCES

A˚ kesson B. 1962. The embryology of Tomopteris helgolandica Hanstro¨m B. 1928. Weitere Beitra¨ge zur Kenntnis des Gehirn (Polychaeta). Acta Zoologica 43:135–200. und der Sinnesorgane der Polycha¨ten (Polygordius, Briggs DEG, Clarkson ENK. 1987. The first tomopterid, a Tomopteris, Scolecolepis). Zeitschrift fu¨r Morphologie polychaete from the Carboniferous of Scotland. Lethaia und O¨ kologie der Tiere 13:329–358. 20:257–262. Kotikova EA. 1973. New data concerning the nervous system Bubko OV. 1981. The nervous system of Protodrilus of Archiannelida. Zoologichesky Zhurnal 52:1611– (Archiannelida). Zoologichesky Zhurnal 60:1749–1755 1615 [in Russian]. [in Russian]. Lagutenko YuP. 1986. Neuropil structure of the abdominal Bubko OV. 1984. The morphology of the nervous system trunk in the Oweniidae (Polychaeta). Arhiv anatomii, in Nerilla antennata (Archiannelida). Zoologichesky gistologii i embriologii 90(2):25–33 [in Russian]. Zhurnal 63:165–170 [in Russian]. Levenstein RY. 1981. Some peculiarities of the distribution of Bubko OV, Minichev YuS. 1972. Nervous system in Owenii- the polychaetous (Fam. Polynoidae) in the Canada dae. Zoologichesky Zhurnal 51:1288–1299 [in Russian]. Basin on the Arctic Ocean. Trudy Instituta Okeanologii Buzhinskaja GN. 2001a. The finding of pelagic polychaetes im. P.P. Sirsova 115:26–36 [in Russian]. of the family Yndolaciidae Støp-Bowitz, 1987 in the Pereyaslawzewa SM. 1896. Me´moire sur l’organisation de la Arctic Ocean. To the question on taxonomic status of Nerilla antennata O. Schmidt. Annales des Sciences the family. In: Otchetnaya nauchnaya sessiya po itogam Naturelles. Zoologie 1:277–345. rabot 2000 g.Tezisy dokladov. 3-5 aprelya 2001 Pleijel F. 1991. Phylogeny and classification of the Phyllodo- Rossiyskaya Akademiya nauk, Zoologichesky Institut, cidae (Polychaeta). Zoologica Scripta 20:225–261. St. Petersburg. p 13 [in Russian]. Buzhinskaja GN. 2001b. Polychaeta. List of species of free Sirenko BI, Marhaseva EL, Buzhinskaja GN, Golikov AA, living invertebrates of Eurasian arctic seas and adjacent Menshutkina TV, Petryashov VV, Semenova TN, deep waters Explorations of the Fauna of the Seas Stepanjants SD, Vassilenko SV. 1996. Preliminary data 51(59):52–66. on suprabenthic invertebrates collected during the RV Chambers SJ, Muir AI. 1997. Polychaeta: British Chryso- Polarstern cruise in the Laptev Sea. Polar Biology petalidea, Pisionoidea and Aphroditoidea. Synopses of 16:345–352. the British Fauna (New Series) 54:1–202. Støp-Bowitz C. 1987. A new genus and species (Yndolacia Fauchald K, Rouse GW. 1997. Polychaete systematics: past lopadorrhynchoides) of pelagic polychaetes, represen- and present. Zoologica Scripta 26:71–138. tative of a new family, Yndolaciidae. Bulletin of the Fraipont J. 1884. Recherches sur la syste´me nerveux central et Biological Society of Washington 7:128–130. pe´riphe´rique des Archiannelides (Protodrilus et Poly- Støp-Bowitz C. 1992. Polyche`tes pelagique des campagnes de gordius)etduSaccocirrus papillocercus. Archives de “L’ Ombango” dans les eaux equatoriales et tropicales Biologie 5:243–304. Ouest-Africaines. Paris: ORSTOM editions.

Accepted 28 June 2004 – Printed 1 October 2004 Editorial responsibility: Egil Karlsbakk Downloaded by [TOBB Ekonomi Ve Teknoloji] at 13:30 21 December 2014