versión impresa ISSN 0716-078X
Rev. chil. hist. nat. v.73 n.4 Santiago dic. 2000
Family Pilargidae (Polychaeta): new distributional ranges and a new record for the Chilean coast
La familia Pilargidae (Polychaeta): nuevos rangos de distribución y un nuevo registro para la costa de Chile
1 Departamento de Ecología, Facultad de Ciencias Biológicas, Pontificia Universidad Católica de Chile,
Casilla 114-D, Santiago, Chile, e-mail: firstname.lastname@example.org
2 Universidad Arturo Prat, Departamento de Ciencias del Mar, Casilla 121, Iquique, Chile
Species of the family Pilargidae are revised based on bibliographic information and on specimens collected in soft bottoms, from 16 to 68 m depth, in three localities of the northern Chilean coast. Five genera and six especies, including a new record for the Chilean fauna, Parandalia fauveli, are recognized. Synonyms, diagnostic features, geographic distribution data and illustrations are provided for each species. Abundance data and biomass, as well as granulometry and organic matter of the sediments where the specimens were collected, are also reported. Presence of Pilargis berkeleyae and Sigambra bassi in the collected material allows extension of their distributional ranges from approximately 35-36ºS to 18º30'S. A taxonomic key to the six species of Pilargidae from the Chilean coast is provided.
Key words: Pilargidae, Chilean coast, taxonomy, geographical distribution, new record.
Se revisan las especies de Pilargidae de la costa de Chile a partir de especímenes recolectados en fondos blandos sublitorales, entre 16 y 68 m de profundidad, en tres localidades del extremo norte del país, y se complementa con información bibliográfica. Se reconocen 5 géneros y seis especies incluyendo un nuevo registro para la fauna de Chile, Parandalia fauveli. Se incluye sinonimia, caracteres diagnósticos, distribución geográfica e ilustraciones para cada especie. De las especies recolectadas se entregan, además, valores de abundancia y biomasa así como también valores de granulometría y contenido de materia orgánica de los sedimentos que habitan. La presencia de Pilargis berkeleyae y Sigambra bassi, en el material recolectado, permite extender sus rangos de distribución hacia el norte de Chile desde aproximadamente 35-36ºS hasta 18º30'S. Se incluye una clave taxonómica para las seis especies de pilárgidos de la costa de Chile.
Palabras clave: Pilargidae, costa de Chile, taxonomía, distribución geográfica, nuevo registro.
The Pilargidae Saint-Joseph has been characterized by some authors as a badly defined taxon whose members, on occasion, have been placed among the Hesionidae or the Syllidae (Hartman 1947, Pettibone 1963). It has also been considered an enigmatic family because its species are, in general, scarce and of a supposedly very ample distribution (Salazar-Vallejo & Orensanz 1991). The pilargid species live in sand or muddy sand, from the intertidal zone down to great depths (Pettibone 1982). Day (1967) considers them as carnivores or omnivores, although according to Fauchald & Jumars (1979) there is no empirical evidence on their feeding habits. Speculations about their feeding habits have been based only on anatomical observations (Wolf 1984).
On a global level, the family includes 50 species distributed in 10 genera (Pettibone 1982). Along the coasts of Chile, the knowledge on the pilargids goes back to findings from the "Lund University Chile Expedition" in 1948-1949 (Wesenberg-Lund 1962), the oceanographic expedition "Marchile I" in 1960, and collections realized in northern Chile (Hartmann-Schröder 1965), the expedition of the USNS "Eltanin" in 1962-1966 (Hartman 1967), and subtidal samples obtained in the continental shelf off Bahía de Concepción (Cañete et al. 1990).
The objective of this study is to contribute to the knowledge of the pilargid species present along the Chilean coast. Based on specimens collected in northern Chile between Arica and Bahía de Mejillones, Antofagasta, diagnostic features, synonyms, geographical distribution and illustrations of each of the 6 species found in Chile are presented. Moreover, abundance and biomass data for the species collected in the present study, as well as granulometry and organic matter content of the sediments in which they were found, are reported.
MATERIALS AND METHODS
The species were collected between Arica (18º30'S) and Bahía de Mejillones (23º05'S), 16-68 m depth, between May 1996 and March 1998, as part of a project on diversity and polychaete abundance in subtidal soft-bottoms in northern Chile.
Three samples of sediment were taken with a 0.1 m2 Van Veen type dredge at each site. The first two were used in obtaining the fauna, which was separated from the sediment with a 1.0 mm mesh net and fixed in 10% formalin neutralized with borax. Specimens were then sorted and identified to the species level, counted and weighed (wet weight, 0.001 g precision scale). The last sample dredged was used in determining the organic matter content and granulometric characteristics of the sediment.
Figures were prepared by means of a camera lucida on a Wild M-5 stereoscopic microscope.
Numbers in parenthesis in the section corresponding to the material examined indicate the number of specimens examined.
The specimens are deposited in the polychaete collection in the Sala de Sistemática, Departamento de Ecología, Pontificia Universidad Católica de Chile, Santiago (SSUC).
KEY TO THE GENERA AND SPECIES OF PILARGIDAE RECORDED
FROM THE SOUTHEASTERN PACIFIC OCEAN OFF CHILE
1 (2) Notopodia with stout protruding hooks or spines ........................3
2 (1) Notopodia without protruding hooks or spines............................ genus Pilargis ..........................Pilargis berkeleyae
3 (4) Notopodia with recurved protruding hooks..................................5
4 (3) Notopodia with stout, straight spines............................................9
5 (6) Antennae shorter than palps; integument papillated................. genus Ancistrosyllis..............................7
6 (5) Antennae longer than palps; integument smooth...................... genus Sigambra....................................Sigambra bassi
7 (8) Ventral cirri beginning on setiger 1; 2 pairs of tentacular cirri..............................................Ancistrosyllis groenlandica
8 (7) Ventral cirri beginning on setiger 4; 1 pair of tentacular cirri Ancistrosyllis quellina
9 (10) Prostomial antennae and peristomial cirri present....................genus Synelmis.................................... Synelmis albini
10 (9) Prostomial antennae and peristomial cirri absent.................... genus Parandalia.............................Parandalia fauveli
Ancistrosyllis groenlandica McIntosh, 1879: 502, pl. 65, figs. 3, 20; Pettibone, 1966: 166-168; Hartman, 1967: 53; Imajima, 1987: 153-155, fig. 2 a-k; Salazar-Vallejo & Orensanz, 1991: 271-272, fig. 2 A-D.
Description and figures adapted from Pettibone (1966).
Description: Up to 40 mm in length, width 1mm, up to 70 segments. Body flattened dorsoventrally, widest in middle region, tapering anteriorly and posteriorly. With conspicuous rust-colored glandular areas in anterior part of notopodial lobe, beginning on setiger 4, and ventrally near bases of parapodia. Integument covered with numerous scattered short papillae. Prostomium (Fig. 1A) with large palpophores and small button-like palpostyles; with three small subequal antennae shorter than palps; without eyes. Proboscis short, cylindrical, with few scattered papillae. Peristomium with two pairs of short subequal cirri, similar to antennae. Dorsal cirri of first segment about twice as long as those of the second segment. Dorsal cirri short and fusiform, extending slightly beyond neuropodial lobes. From setiger 4 to 6, notopodia (Fig. 1B) with somewhat inflated notoacicular lobes and protruding hooked setae (Fig. 1C). Neuropodial lobe conical, with neuroaciculum and neurosetae (Fig. 1C) variable in length. Pygidium rounded, with paired short anal cirri similar to dorsal cirri.
Fig. 1. Ancistrosyllis groenlandica (after Pettibone 1966), A) anterior end in dorsal view; B) middle parapodium; C) notopodial hooked seta and neurosetae. Ancistrosyllis quellina (after Wesenberg-Lund 1962), D) anterior end in dorsal view; E) 16th parapodium from the right side, dorsal view; F) seta. Parandalia fauveli, G) anterior end in dorsal view; H) 12th parapodium; I) neuropodial seta from 12th parapodium.
Distribution: West Greenland; British Isles; Gulf of St. Lawrence to offshore New Jersey; Mediterranean; West Africa; Japan; off northeastern South America (Pettibone 1966, Imajima 1987); Falkland Islands (Hartman 1967); southwestern Chile off Arauco, 957 m, off Isla de Chiloé, 3651-3655 m; Strait of Magellan, Paso del Hambre, 485 m; Drake Passage, NW of South Shetland Islands, 3867-4086 m (Hartman 1967, Rozbaczylo 1985).
Ancistrosyllis quellina Wesenberg-Lund, 1962: 68-73, figs. 22-26.
Description and figures adapted from Wesenberg-Lund (op. cit.).
Description: Complete specimens with 28 and 43 setigers measuring 5 and 9.5 mm, respectively. The colour is yellowish-brown; on the anterior border of each parapodium there is a rusty red patch. Prostomial lobe small and inconspicuous (Fig. 1D); palpophores massive and broad forming the convex margin of the head; they are nearly fused, and from their ventral side a pair of palpostyles arise. Two oblong dark eyespots on each side of the head. A small, unpaired, conical antenna between the eyes; no lateral antennae were observed. Pharynx muscular, cylindrical, very long, reaching backwards, internally, to the 6th setiger; a circle of about 14 large liguliform papillae bordering the oral aperture, was observed through the transparent body; without jaws. Peristomium asetigerous, with a single pair of small, subulate tentacular cirri. The three anteriormost parapodia uniramous, the rest sesquiramous; the former are rather blunt, the rest gradually getting longer and more conical. First parapodium with dorsal cirrus twice as long as those of following parapodia; no ventral cirri in the first three parapodia. From the fourth setiger there is a reduced, liguliform notopodium and a much longer, pointed neuropodium, both with short subulate cirrus. Notopodia (Fig. 1E), from the 4 th segment to the last but two, with a single, large, coarse, strongly curved, transparent hook, rising above the base of the dorsal cirrus. Up to 12 neurosetae, smooth (Fig. 1F). Pygidium with two short, digitiform anal cirri.
Distribution: Up until now, known only for type locality, Golfo de Ancud, SW of Isla Queullín, 300 m.
Remarks: According to Wesenberg-Lund (op. cit.), A. quellina is most similar to A. groenlandica McIntosh, but they differ by the following characteristics present in the former species: only one pair of peristomial cirri; no ventral cirrus in the first three setigers; notopodium with only a single hook (A. groenlandica also has a slender bristle); neuropodial bristles differing in length and without wings or claw. It also has been pointed out the absence of prostomial paired antennae in A. quellina, but this is a characteristic that needs confirmation.
Loandalia americana Hartman, 1947: 506-509, pl. 63, figs. 1-7; 1951: 39, pl. 12, figs. 1-3.
Occurrence: AS004 (2), AS007(2), AS008 (1), MS019 (5)
Description: Length of the larger complete specimen 32 mm, width 1.5 mm, 96 setigers. Those preserved in alcohol are iridescent, light brown in colour. Body elongated, subcylindrical, with inflated anterior region through first 6 setigers (Fig. 1G). Prostomium small, subtriangular, widest at its posterior margin, with palpophores indistinct from rest of prostomium; without antennae, eyes or other visible structures. Palpi biarticulate, directed forward, with minute palpostyles, more or less telescoped within palpophores. Peristomium somewhat longer than the next segment (first setigerous segment); shallowly emarginate at its anterior dorsal edge where it meets the prostomium. First pair of parapodia uniramous, the remainder subbiramous. Notopodia beginning on setiger 2, shaped like small papilar lobes (Fig. 1H), with stout crystal-clear notoacicula present from setiger 7 to the end of body; with additional slender notosetae, 0-3 in number. Neuropodia cylindro-conical with small lobe ("ventral cirrus") on ventral side of their distal ends, from setiger 6. Neurosetae simple (Fig. 1I), geniculate or curved, tapering to fine tips, with numerous transverse spinous rows. Pygidium a rounded anal plate, concave ventrally, with pair of lateral cirri and midventral one; the lateral are much longer and measure approximately 1.5 to 2 times the length of pygidium.
Remarks: This represents the first record of the species in the southeastern Pacific Ocean off Chile.
Distribution: Southern California; Gulf of Mexico (Florida, Mississippi, Louisiana); Central America, Guatemala (Pettibone 1966); Bay of Ancón, Perú (Tarazona et al. 1988, 1996); Chile, from Arica (18º30'S) to Mejillones, Antofagasta (23º05'S), between 16 and 40 m depth.
Ecological remarks: Organisms of this species were collected in Arica, Iquique and Mejillones, in 16 and 40 m depth, in fine sandy-type sediment (1.87 < Ø > 3.14), with low values of mud-clay (0.08 and 4.73%) and total organic matter values in the sediment of 1.71 and 8.88%. Specimens were collected in March 1997 and March 1998 with abundances of 1 and 11 ind/0.2m2, and biomass ranging from 0.01 to 6.02 g/0.2m2.
Pilargis berkeleyae: Pettibone, 1966: 161-164, figs. 1-2; Kirkegaard, 1983: 211; Imajima, 1987: 162-164; Cañete, Gallardo & Carrasco, 1990: 154-156; Salazar-Vallejo & Orensanz, 1991: 274-275, fig. 3 a-c.
Occurrence: MS013 (1), MS006 (1)
Description: Length of the largest complete specimen 23 mm, width 2.5 mm, 97 setigers. Colour in preserved specimens yellowish-brown. Body flattened, ribbon-like, tapered anteriorly and posteriorly. Integument minutely papillated on body and appendages. Prostomium small (Fig. 2A) with paired short lateral antennae; with ovate palpophores and minute palpostyles. Peristomium fused with prostomium, with two pairs of short tentacular cirri, fusiform, dorsal pair slightly longer than ventral pair. All parapodia subbiramous (Fig. 2B); notoacicula colorless, without notosetae; dorsal cirri conical to ovate. Dorsal cirri of first setigerous segment longer than following. From setiger 7 notopodia with cirrophores clearly thicker than those of anteriormost segments. Reddish to dark-brown glandular areas in anterior face of notopodial cirrophores from setiger 9-10 to the end of the body; also with small pigmented glandular areas at the ventral base of neuropodia from setiger 10 to approximately the end of body. Neuropodia conical, with stout neuroacicula and bundle of simple neurosetae; neurosetae variable in length, finely serrated on anterior edge, falcate and bidentate; ventral cirri conical. Pygidium rounded with 2 conical anal cirri.
Fig. 2. Pilargis berkeleyae, A) anterior end in dorsal view; B) parapodium 20. Sigambra bassi, C) anterior end in dorsal view; D) parapodium 20. Synelmis albini (after Hartmann-Schröder 1965), E) anterior end in dorsal view; F) posterior parapodium in posterior view; G) inferior neuropodial seta and forked neuroseta.
Distribution: Washington to southern California (Pettibone 1966); Posyet Bay (Sea of Japan); Japan (Imajima 1987); West Africa; Liberia, off the Congo River; Angola (Kirkegaard 1983); Río de la Plata, off Uruguay (Salazar-Vallejo & Orensanz 1991); Chile, from Arica (18º30' S) to Concepción (36º31'S), between 23 and 70 m depth.
Ecological remarks: Organisms of this species were collected in Arica, Iquique and Mejillones in 23 and 68 m depth, in fine sandy-type sediments (2.64 < Ø > 3.14), with low values of mud-clay (0.33 and 4.73%) and total organic matter values in the sediments of 1.71 and 2.15%. Specimens were collected in November 1997 and March 1998 with abundances of 1 and 9 ind/0.2m2 and biomass ranging from 0.05 to 0.12 g/0.2m2
Occurrence: AS001(1), TS002(1), TS012(1), MS004(2)
Description: Length of the largest complete specimen 20 mm, width 2 mm, 105 setigers. Colour in preserved specimens yellowish-brown. Integument smooth, without papillae. Body flattened dorsoventrally, widest in anterior region. Prostomium (Fig. 2C) with biarticulate palps with large palpophores and small palpostyles; with a median antenna on posterior part of prostomium and two shorter ones laterally. There are no eyes. The everted proboscis is cylindrical, with circlet of 14 conical papillae distally. Peristomium, or tentacular segment, longer than following ones, with 2 pairs of tentacular cirri similar to lateral antennae. First setigerous segment with a pair of dorsal cirri similar to the median prostomial antenna; the ventral cirri are much smaller, resembling those farther back. All parapodia subbiramous (Fig. 2D); notopodia low conical, with stout notopodial hooked notoseta beginning on about setiger 15; dorsal cirri long, subulate. Neuropodia conical, with neuroaciculum and numerous capillary neurosetae of different length; longer ones slender, smooth or faintly serrated, shorter ones serrated; ventral cirri cirriform. Pygidium with paired long anal cirri.
Distribution: North Carolina; Florida; central and southern California, intertidal to 33 m (Pettibone 1966, Gardiner 1975); Chile, from Arica (18º30'S) to Concepción (36º32'S), between 19 and 94 m depth.
Ecological remarks: This species was collected in Arica, Tocopilla and Mejillones, in 19 and 68 m depth, in fine and very fine sandy-type sediments (1.87 < Ø > 3.14), with low values of mud-clay (0.08 and 4.73%) and total organic matter values in the sediments of 1.71 and 8.88%. Specimens were collected in November 1997 and March 1998 with abundances of 1 and 2 ind/0.2m2 and biomass ranging from 0.01 to 0.04 g/0.2m2.
Ancistrosyllis albini Langerhans, 1881: 107-108, fig. 16 a-e.
Ancistrosyllis rigida: Hartmann-Schröder, 1965: 285-287, figs. 286-288.
Synelmis albini: Pettibone, 1966: 191-195, figs. 19-21; Westheide, 1974: 223-225, figs. 12-13;
Description: Length to 60 mm, width to 1.5 mm, up to 170 segments. Body elongate, subcylindrical, tapered posteriorly, segmental grooves indistinct anterodorsally, with parapodia distinctly set off from body. Colour in preserved specimens yellowish to reddish, dark brown, iridescent, with reddish glandular areas at bases of parapodia. Integument with thick, smooth, iridescent cuticle, without papillae. Prostomium with pair of biarticulate palps sometimes turned ventrally; palpophores large, oval; palpostyles small, oval, sometimes retracted within palpophores; with 3 short antennae; with pair of posterior eyes, each 2-4 contiguous spots. Proboscis long, cylindrical, muscular, without papillae. Peristomium (tentacular segment) prominent, overhanging prostomium and forming ventral lip, achaetous, with 2 pairs of subequal tentacular cirri. Dorsal cirri of first setiger similar to following. Parapodia subbiramous, with dorsal and ventral cirri subequal, subulate to oval with acuminate tips. Notopodia with 1-2 notoacicula and stout, protruding, straight or slightly bent, acicular notosetae, beginning on setiger 5-20 (difficult to detect, not emergent at first). Neuropodia short, cylindrical, with 3 neuroacicula and bundle of neurosetae. Neurosetae limbate, tapering to capillary tips. Pygidium rounded, with pair of anal cirri similar to dorsal and ventral cirri.
Remarks: This species is known in Chile only through its record in Taltal (25º25'S), by Hartmann-Schröder (1965).
Distribution: Widespread in tropical and subtropical waters; Canary Islands; southern to Lower California; Panama; Florida; Marshall Islands; Tuamotu Islands; Gambier Islands; Red Sea; Indian Ocean; Japan (Imajima 1987); North Carolina to Brazil (Gardiner 1975); Galapagos Islands (Westheide 1974); Chile: Taltal, 0.5 m (Hartmann-Schröder 1965).
In the southeastern Pacific Ocean, off the Chilean coast, a total of 6 pilargid species distributed in 5 genera have been registered until now. The distribution ranges are extended farther north for two of the 5 previously known species (Pilargis berkeleyae Monro, 1933 and Sigambra bassi (Hartman, 1947)) and Parandalia fauveli (Berkeley and Berkeley, 1941) is registered for the first time in the southeastern Pacific Ocean between Arica and Antofagasta. These results support the hypothesis that a lower number of registered polychaete species in northern Chile, between 18º and 32ºS, in comparison to central (32º-44ºS) and southern (44º-56ºS) Chile is due, in part, to low sampling intensity in this zone, compared to the other two (Rozbaczylo & Simonetti, 2000).
The first bioecological antecedents on the pilargid species on the coast of Chile are given for posterior comparisons with other localities. The species Parandalia fauveli, Pilargis berkeleyae and Sigambra bassi are characterized by inhabiting fine sand sediments, with low mud-clay and total organic matter values between 1.71 and 8.88%; the lowest abundance values were found in Sigambra bassi (1-2 ind/0.2 m2), while the highest reached abundance was for Parandalia fauveli (11 ind/0.2m2).
We are grateful to the group at the Benthos Invertebrate and Zoology Laboratory of Universidad Arturo Prat for their support in the field and the collection of samples. We thank Clara Yañez for her patience and dedication in the preparation of the drawings. The first author thanks the editors Juan Carlos Castilla and Bernabé Santelices for their kind invitation to participate in this special volume dedicated to the memory of our dearly remembered Professor Patricio Sánchez R.
BERKELEY E & C BERKELEY (1941) On a collection of Polychaeta from southern California. Bulletin of the Southern California Academy of Sciences 40: 16-60. [ Links ]
CAÑETE JI, VA GALLARDO & FD CARRASCO (1990) On the Presence of Pilargis berkeleyae Monro, 1933 (Polychaeta, Pilargidae) in the soft bottoms of central Chile shelf. Boletín de la Sociedad de Biología de Concepción (Chile) 61: 153-156. [ Links ]
DAY JH (1967) A Monograph on the Polychaeta of Southern Africa, Part I: Errantia. Publications of the British Museum (Natural History) (London) 656: 1-458. [ Links ]
EMERSON RR & K FAUCHALD (1971) A revision of the genus Loandalia Monro with description of a new genus and species of pilargiid polychaete. Bulletin of the Southern California Academy of Sciences 70: 18-22. [ Links ]
FAUCHALD K & PA JUMARS (1979) The diet of worms: a study of polychaete feeding guilds. Oceanography and Marine Biology Annual Review 17: 193-284. [ Links ]
GARDINER SL (1975) Errant polychaete annelids from North Carolina. Journal of the Elisha Mitchell Scientific Society 91: 77-220. [ Links ]
HARTMAN O (1947) Polychaetous annelids. Part VIII. Pilargiidae. Allan Hancock Pacific Expedition 10: 483-523. [ Links ]
HARTMAN O (1951) The littoral marine annelids of the Gulf of Mexico. Publications of the Institute of Marine Science 2: 7-124. [ Links ]
HARTMAN O (1967) Polychaetous annelids collected by the USNS Eltanin and Staten Island Cruises, chiefly from Antarctic Seas. Allan Hancock Monographs in Marine Biology 2: 1 - 387. [ Links ]
HARTMAN O (1968) Atlas of the Errantiate Polychaetous Annelids from California. Los Angeles, California. University of Southern California, Allan Hancock Foundation. 828 pp. [ Links ]
HARTMANN-SCHRÖDER G (1965) Zur Kenntnis des Sublitorals der chilenischen Küste unter besonderer Berücksichtigung der Polychaeten und Ostracoden. Tl. II. Die Polychaeten des Sublitorlas. Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut 62 (Supplement): 59 - 305. [ Links ]
IMAJIMA M (1987) Pilargidae (Annelida, Polychaeta) from Japan. Bulletin of the National Science Museum, Tokyo, Series A (Zoology), 13: 151 - 164. [ Links ]
KIRKEGAARD JB (1983) The polychaeta of West Africa. Part II. Errant species. 1. Aphroditidae to Nereididae. Atlantide Report 13: 181 - 240. [ Links ]
LANGERHANS P (1881) Über einige canarische Anneliden. Nova Acta Leopoldina, Königliche Deutschen Akademie der Naturforscher 42: 93 - 124. [ Links ]
McINTOSH WC (1879) On the Annelida obtained during the cruise of H.M.S. Valorous to Davis Strait in 1875. Transactions of the Linnean Society of London, series 2, 1: 499-511. [ Links ]
MONRO CCA (1933) On a new species of polychaeta of the genus Pilargis from Friday Harbour, Washington. Annals and Magazine of Natural History, series 10, 11: 673-675. [ Links ]
PETTIBONE MH (1963) Marine polychaete worms of the New England region, 1: Families Aphroditidae through Trochochaetidae. United States National Museum Bulletin 227: 1 - 356. [ Links ]
PETTIBONE MH (1966) Revision of the Pilargidae (Annelida: Polychaeta), incluiding descriptions of a new species and descriptions of the pelagic Podarmus ploa Chamberlin (Polynoidae). Proceedings of the United States National Museum 118: 155 - 207. [ Links ]
PETTIBONE MH (1982) Annelida. In: Parker SB (ed) Synopsis and Classification of Living Organisms 2: 1-43. New York, McGraw Hill. [ Links ]
ROZBACZYLO N (1985) Los Anélidos Poliquetos de Chile. Indice sinonímico y distribución geográfica de especies. Monografías Biológicas (Chile) 3: 1-284. [ Links ]
ROZBACZYLO N & JA SIMONETTI (2000) Diversity and distribution of Chilean benthic marine polychaetes: state-of-the art. Bulletin of Marine Science 67: 1 - 14. [ Links ]
SALAZAR-VALLEJO SI & JM ORENSANZ (1991) Pilárgidos (Annelida: Polychaeta) de Uruguay y Argentina. Cahiers de Biologie Marine 32: 267-279. [ Links ]
TARAZONA J, H SALZWEDEL & W ARNTZ (1988) Oscillations of macrobenthos in shallow waters of the Peruvian central coast induced by El Niño 1982-83. Journal of Marine Research 46: 593-611. [ Links ]
TARAZONA J, W ARNTZ & E CANAHUIRE (1996) Impact of two "El Niño" events of different intensity on the hypoxic soft bottom macrobenthos off the central Peruvian coast. P.S.Z.N. I: Marine Ecology 17: 425 - 446. [ Links ]
WESENBERG-LUND E (1962) Reports of the Lund University Chile Expedition 1948-49, 43: Polychaeta Errantia. Lunds Universitets Årsskrift, new series, section 2, 57: 1-137. [ Links ]
WESTHEIDE W (1974) Interstitielle fauna von Galapagos XI. Pisionidae, Hesionidae, Pilargidae, Syllidae (Polychaeta). Mikrofauna des Meeresbodens 44: 1-146. [ Links ]
WOLF PS (1984) Family Pilargidae. In: Uebelacker JM & PG Johnson (eds) Taxonomic Guide to the Polychaetes of the Northern Gulf of Mexico 4: 29.1-29.41. Mobile, Alabama: Barry A Vittor and Associates. [ Links ]
Invited Editor J.C. Castilla