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Revista chilena de historia natural

versión impresa ISSN 0716-078X

Rev. chil. hist. nat. v.74 n.3 Santiago set. 2001 

Polychaetes from a southwestern shallow shelf Atlantic area
(Argentina, 38 S) affected by sewage discharge

Poliquetos de una plataforma somera del Atlántico sudoccidental
(Argentina, 38 S) afectada por efluentes cloacales


1Departamento de Ciencias Marinas, Universidad Nacional de Mar del Plata, Deán Funes 3350 (7600), Mar del Plata, Argentina, e-mail:
2Consejo Nacional de Investigaciones Científicas y Tecnológicas & Instituto Nacional de Investigación y Desarrollo Pesquero, Casilla de Correos 175 (7600) Mar del Plata, Argentina


A benthos-sampling program was carried out in the coastal area of Mar del Plata city in order to know the composition and distribution of infaunal polychaetes living on a subtidal soft-bottom substrate affected by the sewage discharge. Sandy bottoms showed heterogeneous sediment composition and very low organic matter content. Thirty three polychaete species were identified in a range depth between 6.5 to 12.5 m. A comparison with data from 30 years ago from adjacent areas deeper than 12 m is herein presented. Polychaete distribution was related both to the type of sediment and to the content of organic matter. Owenia fusiformis Delle Chiaje was dominant and peaked at stations near the sewage effluent. Small-sized polychaetes were abundant in the sampling area. A possible explanation to the faunal composition of sites near sewage effluents is that hydrodynamic forces frequently disturb the area, and benthic fauna is permanently in an early stage of development. The effect of sewage appeared to be restricted to the shallower subtidal area around the outfall.

Key words: polychaetes, domestic outfall, sandy subtidal bottoms, South Atlantic.


Un programa de muestreo bentónico fue llevado a cabo en la ciudad de Mar del Plata con el fin de examinar la composición y distribución de los poliquetos infaunales de los sustratos submareales afectados por el vertido cloacal. Los fondos arenosos muestran heterogeneidad en su composición de sedimentos y un bajo contenido de materia orgánica. Se identificó un total de 33 especies de poliquetos en el rango de profundidades comprendido entre 6,5 y 12,5 m. En este trabajo se presenta una comparación de estos datos con datos de 30 años atrás provenientes de fondos de más de 12 m de profundidad. El patrón de distribución de los poliquetos se relaciona tanto con el tipo sedimentario como con el contenido de materia orgánica. Owenia fusiformis Delle Chiaje fue la especie dominante en las estaciones más cercanas al efluente. Dicha área estuvo caracterizada por la presencia de poliquetos de tamaño pequeño. Una explicación probable para la composición faunística en estos ambientes bentónicos cercanos a descargas es que los procesos hidrodinámicos perturban frecuentemente el área, y la fauna bentónica se encontraría permanentemente en un estado de desarrollo inicial. El efecto del efluente cloacal parece estar limitado al área submareal somera alrededor de la descarga.

Palabras clave: poliquetos, efluente cloacal, fondos submareales arenosos, Atlántico sur.


Organic enrichment of sediments is perhaps the best-documented disturbance on marine environments (Pearson & Rosenberg 1978). To assess the impact of man-made changes, it is useful the use of benthic fauna, at the level of species, population or community.

One of the most neglected, major group of marine invertebrates may be the polychaetous Annelids that could be useful as indicators of varying degrees of marine pollution (Reish 1980). Polychaetes are used as monitors for toxic materials, as bioassay organisms, and as pollution indicators at the various levels for marine environmental quality (Pocklington & Wells 1992). Reish (1955) has early mentioned the relationship between Polychaetous Annelids and harbor pollution. After this, many papers have used these organisms in qualitative and/or quantitative studies about pollution (e.g., Reish 1957, 1963), including an Annelid Index of Pollution (Bellan 1980) to assess pollution at a site affected by municipal sewage (Bellan et al. 1988). Some species of the families Capitellidae, in particular Capitella capitata (Fabricius 1780), and Spionidae have become widely accepted as organic pollution indicators (Tsutsumi 1990, Pocklington & Wells 1992).

In Argentina, studies on the relationship between polychaetes and pollution are scarce, and are related to the community level. In Blanca bay (39 S), a mixohaline shallow environment, the presence of an infaunal community dominated by Polychaetes was found in relation to organic enrichment (Elías 1992, Elías & Bremec 1995).

The Mar del Plata city has only a pre-treatment plant for sewage of 800,000 inhabitants, and the effluent discharges directly on an intertidal northern beach, with a continuous rate flow between 2.5 to 8.0 m3 seg-1 (in rainy days). However, the city is the greatest recreational place of Argentina, and receives more than 2,000,000 people during summer. The city also has a port that concentrates the catches and commercialization of coastal fisheries. Municipal government has the intention to construct a submarine pipe 3 km seaward, where sand bottoms are at 11 m depth. Previous benthic studies in the area of Mar del Plata were carried out in bottoms deeper than 12 m, almost 30 years ago (Olivier et al. 1968). Very scarce information was obtained till present about benthic communities of the shallow soft-bottoms.

A monitoring program was carried out in this area to know the composition and distribution of macrozoobenthos, to have reference data for comparing the present condition with the resulting changes during and after the pipe construction. In this contribution we describe the polychaete composition and distribution in subtidal bottoms between 6.5 to 12.5 m depth, in front of the domestic sewage discharge. The aim of this study is to detect a polychaete distribution pattern useful for assessing organic enrichment in the area.


Study area

The sampling area is an open coast subjected to the littoral current (south to north) and to autumn-winter storms (predominantly south-southeast).

Sand ridges dominate the northern areas, while sand ribbons characterize the southern part. Ridges are composed by fine sand with coarse material (mainly shells) in the swales, while sand ribbons are fine-sand bodies (Isla & Schnack 19861). The sewage flume is mostly directed to north, but with strong north winds it falls to the south (Isla & Ferrante 1997). Organic matter values in sediment range between 0.21 to 0.76 % (Isla et al. 1997).

Sampling procedures and analysis

During 1996 we collected 12 samples from subtidal bottoms (between 6.5 to 12.5 m depth) using a benthic dredge (600 x 250 x 200 mm) during 10 min each (Fig. 1). The sediment was sieved (1 mm) and the retained polychaetes were identified and counted. Sediment analysis (by dry sieving) was performed for subsamples of sediments, including the determination of grain-size parameters in phi units (-log2 of particle diameter) and organic matter content (Isla et al. 1997). Polychaete species were ranked according to their dominance (number of individuals/total number 100). The frequency (%) was also calculated for each species, and the Shannon index of diversity (Shannon & Weaver 1963) was calculated for each sampling unit. Feeding guilds followed Fauchald & Jumars (1979). Classification and ordination were performed with abundance (log10 X + 1) in the polychaete species (22) with frequency up to 20 % by using Cluster (1-Pearson r index, UPGMA), and Multi-dimensional Scaling (MDS) analyses (Clarke & Warwick 19932).

Fig. 1: Location of sampling area and sampling stations around sewage outfall in the shallow shelf area in front of the Mar del Plata city (38° S, Argentina).

Ubicación geográfica del área y de las estaciones de muestreo alrededor del efluente cloacal en la plataforma somera frente a la ciudad de Mar del Plata (38° S, Argentina).


Polychaetes comprised 33 species and 1,356 individuals, being the dominant group in the community (37.93 % of species) and second in total abundance (30.55 %). The most abundant polychaete species in the area were Owenia fusiformis Delle Chiaje 1844 (3rd in the general rank order), followed by Armandia sp., Ampharete sp., Glycera americana Leidy 1855 and Prionospio sp. The most frequent species were Prionospio sp. and Lumbrineris tetraura (Schmarda 1861), followed by G. americana and Ampharete sp., but O. fusiformis, Armandia sp., Aglaophamus uruguayi Hartman 1953, Diopatra viridis Kinberg 1865 and Phyllodoce sp. also reached high frequency (Table 1). All specimens were juveniles or small-sized polychaetes.


Dominance (%) and frequency (%), and feeding guilds of polychaete species found in the shallow shelf area in front of Mar del Plata city affected by domestic sewage. Feeding guilds follow Fauchald & Jumars (1979); the three-letter code denotes feeding type (F for filter-feeders, C for carnivores, S for superficial deposit-feeders, and B for subsuperficial deposit-feeders), motility (M for motile, D for discretely motile, and S for sessile), and feeding structures (J for jawed, P for pumping, T for tentacled, and X for other)

Dominancia (%), frecuencia (%) y gremios tróficos de poliquetos hallados frente a la plataforma somera de la ciudad de Mar del Plata afectada por efluentes cloacales. La asignación de gremios sigue el criterio de Fauchald & Jumars (1979); el código de tres letras indica tipo de alimentación (F para filtradores, C para carnívoros, S para alimentadores de depósito superficial y B para alimentadores de depósito subsuperficial), movilidad (M para móviles, D para discretamente móviles y S para sésiles) y presencia de estructuras alimentarias (J para mandibulados, P por bombeo, T para tentaculados y X para otras estructuras)

Owenia fusiformis reached higher dominance in three stations, two of them closest to the sewage effluent (4 and 65) with abundances up to 400 ind sample-1 and also associated to low values of both species number and diversity (Table 2). In Cluster and MDS (Fig. 2 and 3), these stations were grouped around a mean phi of 3.0 ± 0.41 and they had the highest organic matter content of sediments (between 0.48 and 0.76 %). Diversity, especially near the effluent, was low. The highest dominance at these stations was achieved by Owenia fusiformis, and, secondarily, by Ampharete sp. The Station 63, the richest and most diverse, was dominated by Aglaophamus uruguayi, Glycera Americana, and Prionospio sp. (with moderate dominances between 19 to 14 %), exhibited a phi of 3.17, an organic matter content of 0.48 %, and it was associated in cluster analysis with the Owenia-dominated stations.

Fig. 2: Cluster analysis (1-Pearson r Index, UPGMA) performed on the abundance of 22 polychaete species having a frequency of 20 % or more (data transformed to log10 X + 1), which were found at 12 subtidal stations in front of the Mar del Plata's sewage effluent.

Análisis de agrupamiento (índice 1-Pearson r, UPGMA) sobre la abundancias de 22 especies cuya frecuencia es de 20 % o superior (datos transformados a log10 X + 1), halladas en 12 estaciones submareales frente a la descarga cloacal de la ciudad de Mar del Plata.

Fig. 3: Multi-dimensional scaling analysis of 12 subtidal stations located in front of Mar del Plata sewage, which included 22 polychaete species whose frequency was 20 % or more.

Análisis de escalamiento multidimensional de 12 estaciones submareales frente a la descarga cloacal de Mar del Plata con la inclusión de 22 especies de poliquetos cuya frecuencia fue de 20 % o superior.

Stations 1, 2, 3, 9 and 62 on the one hand, and stations 10 and 11 on the other, were grouped (in opposite sides) along Dimension 1. Former stations were characterized by the dominance of Armandia sp. and Ampharete sp., or by Prionospio sp. with Armandia sp. (or Magelona riojai Jones 1963, in Station 50) around a mean phi of 1.98 ± 0.78, and low organic content (0.21 %). The hard-bottom species of Syllidae or Dodecaceria concharum Saint-Joseph, 1898, respectively, characterized stations 11 and 10.

Graphically, the two major groups were distributed in fringes parallel to coastal line (Fig. 4). Dimension 1 seemed influenced by grain-size parameters.

Fig. 4: Polychaete assemblages defined by cluster and multi-dimensional scaling analyses. Armandia sp. or Prionospio sp. dominance defines group 1, while Owenia fusiformis dominates in shallow stations of group 2.

Asociaciones de poliquetos definidas por análisis de conglomerados y de escalamiento multidimensional. El grupo 1 está definido principalmente por la dominancia de Armandia sp. o de Prionospio sp., mientras que Owenia fisiformis domina en las estaciones someras del grupo 2.


The only available information about polychaetes in the area of Mar del Plata is referred to bottoms deeper than 12 m and is represented by records cited in Olivier et al. (1968), and taxonomic reports by Orensanz (1973, 1974).

The onuphids Onuphis eremita Audouin & Milne Eduards 1833 and O. setosa Kinberg 1865 were found in waters shallower than previous records, more than 40 m in the former, and 13 to 25 m the latter, while Diopatra viridis was frequent in the subtidal sandy bottoms of the area (Orensanz 1974). In Talcahuano bay (Chile), an environment subjected to sewage outfall, Diopatra chilensis Quatrefages 1865 is the third species in terms of biomass (Oyarzún et al. 1987). Two lumbrinerids were recorded, but only Lumbrineris tetraura was the most frequent. This species has been cited for sandy bottoms of the area, being a common predatory polychaete (Orensanz 1973, Carrasco & Oyarzún 1988). Oyarzún et al. (1987) found a very important biomass of L. tetraura in the nearest station in front of the Talcahuano port affected by sludge from factories and municipal sewage. However, it has been suggested that if the genus Lumbrineris is absent, and the diversity is low, the chance that the site is impacted is great (Rygg 1985). Other frequent predatory polychaete species were Glycera americana, common in the area at all depths, Aglaophamus uruguayi, Syllidae, and Phyllodoce sp.

The pattern of polychaete distribution reported herein agrees with previous analyses carried out at a community level in the study area (Elías et al. 19973), and in which sampling stations formed three groups after conducting principal component and cluster analyses. One of them contains the "rich" stations (mean species number S = 34, mean abundance A = 544 ind.sample-1) in fine and very fine sand (phi = 3). An association characterized by juveniles of the bivalve Amiantis purpuratus (Lamarck 1835), tanaidaceans and amphipods inhabits these stations. Other group cluster of "poor" stations (mean S = 19, mean A = 128), located in coarse sand and gravel with shell debris (phi between 1 and 2), was dominated by the crustacean Corophium sp., the polychaetes Armandia sp. and Ampharete sp., and amphipods. A third group, with the high loading stations 4 and 65 (the closest to the effluent), is dominated by the polychaete Owenia fusiformis. The similarity of these results reveals the important role of polychaetes in structuring the subtidal community.

On the other hand, other benthic associations occupied bottoms deeper than 12 m. The shallowest one is characterized by the anthozoan Renilla sp., the echinoderm Encope emarginata (Leske 1778), and the amphioxus Branchiostoma platae Hubbs 1922 (Olivier et al. 1968). Owenia fusiformis was widely distributed in the stations deeper than 12 m, but rare (Olivier et al. 1968). The occurrence of a relatively high and stable population abundance of O. fusiformis (45 % of the total biomass of the assemblage) was also reported in relation to domestic sewage in the Mediterranean Sea, which is supported by the increased possibilities of flow changes at the mouth of the harbor (Pinedo et al. 1997). In the Talcahuano bay, in the vicinity of a sewage discharge, O. collaris (= O. fusiformis) is also frequent (Oyarzún et al. 1987). Owenia fusiformis is a surface-deposit feeder (Fauchald & Jumars 1979, Gambi 1989) that alternates its feeding mechanisms in relation with the environmental conditions. When high planktonic inputs are produced and flow conditions change, O. fusiformis can behave as a filter feeder (Gambi 1989). Desroy et al. (1997) show that filter feeding predominates at the adult population level, while juveniles alternate between deposit and filter feeding activities under still water conditions at similar frequencies. In the Mar del Plata area, the great south to north current with the high suspended matter due to the domestic sewage allows this species to be enhanced at intermediate distances from the effluent.

Capitella capitata, a classical indicator of organically enriched sediments, appeared once and at low abundance. However, it is frequent and abundant in the interstitial sediments of intertidal mussel beds developed around the effluent (Elías et al. 19994).

It is remarkable that the shallower shelf was never studied faunistically, and several species were new records for Argentine waters, like Magelona riojai (Elías & Bremec 1997), and the genus Armandia (Elías & Bremec unpublished results). Besides, at least two species (Aricidea and Prionospio) seem to be either new for the area or new species. The fringe between 6.5 to 12.5 m depths seems characterized by small-sized polychaetes, like Owenia fusiformis, Prionospio sp., Magelona riojai, Scoloplos (S.) sp., Paraprionospio pinnata (Elhers 1901), Armandia sp., and Glycera americana. Most of these polychaete species are deposit feeders or carnivores, preying upon other polychaetes or small invertebrates, while suspensivores are scarce (Table 1).

A similar association of small polychaetes was observed in the northern shallow shelf of Chile and Perú, characterized by species of the genus Aricidea, Tharyx, Prionospio, Owenia, Magelona and Leitoscoloplos, and also by the medium-sized Paraprionospio pinnata (Carrasco 1997). The frequent dominance of the macrobenthos by small-sized forms has been already reported (Beukema 1988). A model proposed by Rumohr et al. (1996) for benthic communities at the Baltic sea predicted a intermediate successional stage (III) dominated by small polychaete worms (or their ecological equivalents, like amphipods) in relation to a gradient of bottom organic enrichment.

Other possible explanation for the presence of small-sized polychaetes is that hydrodynamics forces frequently disturb the area, and benthic fauna is permanently in an early stage of development. Recolonization following complete defaunation due to disturbances caused by winds and storms is frequently reported on benthic environments (see Probert 1984). Small, opportunistic, tube-dwelling polychaetes are the first faunal components to colonize new or newly disturbed bottoms (Rhoads & Boyer 1982). In the shallow shelf of southern Brazil, it is suggested that after the winter defaunation a recolonization period by opportunistic species (like spionid Polychaetes) takes place (Paiva 1993). In Talcahuano bay (Chile), the small opportunistic spionid Cossura chilensis Hartmann-Schröeder, 1965 dominated bottoms affected by domestic sewage (Oyarzún et al. 1987).

Physical disturbances could be responsible for the low diversity, the small size of dominant fauna, and for the great variability in species number and relative abundance of these assemblages. Sedimentary pattern found in this study agrees with a high-energy environment (Isla et al. 1997). Other studies carried out on macrobenthos of the Argentine shelf also revealed a great bottom heterogeneity, even in relatively small areas, and probably due to hydrodynamic forces (Olivier et al. 1968, Roux et al. 1993). These facts suggest that the main structuring factor in the shallow shelf is the physical disturbance due to strong hydrodynamic forces. However, it is suspected that between 0-6 m depth sewage discharge will be also an important structuring factor of both polychaete populations and macrobenthic communities.

1 ISLA FI & EJ SCHNACK (1986) Gradación "ribbons"/dorsales de arena en la plataforma costera entre Mar Chiquita y Mar del Plata. Extended Abstracts from Primera Reunión Argentina de Sedimentología, La Plata, Argentina: 45-48.

2 CLARKE B & R WARWICK (1993) Environmental Effects on Benthic Communities. Lecture Notes for SEAS/EPOS II Workshop, Plymouth Marine Laboratory. 144 pp.

3 ELÍAS R, EA VALLARINO, RO BASTIDA, JP MARTIN & EN IENO (1997) Mar del Plata's sewage impact on coastal soft-bottom benthic communities, Argentina. Extended Abstract of the VII Congreso Latinoamericano de Ciencias del Mar, Santos, Brasil, volume I: 276-278.

4 ELÍAS R, EA VALLARINO, CS BREMEC & C GRAVINA (1999) Los poliquetos de la comunidad intermareal como bioindicadores de enriquecimiento orgánico en el Atlántico Sudoccidental. Extended Abstracts of the VIII Congreso Latinoamericano de Ciencias del Mar, Trujillo, Perú, volume II: 904-905.


To the crew of Coast Guard CG-64 for logistic assistance. The student Julia Desojo and Lic. J.P. Martin participated in sampling and sorting. We are indebted to Dr. F.I. Isla for including us in the sampling program and for advice in sediment analisis. This work was funded by the Obras Sanitarias Sociedad de Estado (Mar del Plata). Stuart Anderson and two anonymous reviewers improved the english version. This paper was read in the Sixth International Polychaete Conference, 2-7 August of 1998, at Curitiba, Brazil. This study corresponds to the scientific contribution 108 from Departamento de Ciencias Marinas, and to the scientific contribution 1147 from Instituto Nacional de Investigación y Desarrollo Pesquero (INIDEP).


BELLAN G (1980) Relation of pollution to rocky substratum polychaetes on the French Mediterranean coast. Marine Pollution Bulletin 11: 318-321.         [ Links ]

BELLAN G, G DESROSIERS & A WILLSIE (1988) Use of an Annelid Pollution Index for monitoring a moderately polluted littoral zone. Marine Pollution Bulletin 19: 662-665.         [ Links ]

BEUKEMA JJ (1988) An evaluation of the ABC-method (abundance/biomass comparison) as applied to macrobenthic communities living on tidal flats in the Dutch Wadden Sea. Marine Biology 99: 425-433.         [ Links ]

CARRASCO FD (1997) Sublittoral macrobenthic fauna off Punta Coloso, Antofagasta, northern Chile: high persistence of the polychaete assemblages. Bulletin of Marine Science 60: 443-459.         [ Links ]

CARRASCO FD & C OYARZÚN (1988) Diet of the polychaete Lumbrineris tetraura (Schmarda) (Lumbrineridae) in a polluted soft-bottom environment. Bulletin of Marine Science 42: 358-365.         [ Links ]

DESROY N, F OLIVIER & C RETIERE (1997) Effects of individual behaviors, inter-individual interactions with the adult Pectinaria koreni and Owenia fusiformis (Annelida, Polychaeta), and hydrodynamism on Pectinaria koreni recruitment. Bulletin of Marine Science 60: 547-558.         [ Links ]

ELÍAS R (1992) Quantitative benthic community structure in Blanca Bay and its relationship with organic enrichment. Publicazione di la Stazione Zoologica di Napoles I, Marine Ecology 13: 189-201.         [ Links ]

ELÍAS R & CS BREMEC (1995) Biomonitoring water quality using benthic communities in Blanca Bay, Argentina. The Science of Total Environment 158: 45-49.         [ Links ]

ELÍAS R & CS BREMEC (1997) First record of Magelonidae, Magelona riojai Jones, 1963 (Polychaeta), in coastal waters of Argentina. Neritica 11: 111-117.         [ Links ]

FAUCHALD K & PA JUMARS (1979) The diet of worms: a study of polychaete feeding guilds. Oceanography and Marine Biology Annual Reviews 17: 193-284.         [ Links ]

GAMBI MC (1989) Osservazioni su morfologia funzionale e comportamento trophico di Owenia fusiformis Delle Chiaje (Polychaeta, Owenidae) in rapporto ai fattori ambientali. Oebalia 15: 145-155.         [ Links ]

ISLA FI & A FERRANTE (1997) Corrientes. In: Isla FI (ed) Estudio del sector de plataforma receptor de la descarga cloacal de Camet, Mar del Plata: 83-116. Final report to Obras Sanitarias Sociedad de Estado, Buenos Aires, Argentina.         [ Links ]

ISLA FI, S SERNA & M FARENGA (1997) Sedimentos del fondo. In: Isla FI (ed) Estudio del sector de plataforma receptor de la descarga cloacal de Camet, Mar del Plata: 70-77. Final report to Obras Sanitarias Sociedad de Estado, Buenos Aires, Argentina.         [ Links ]

OLIVIER SR, R BASTIDA & MR TORTI (1968) Resultados de las campañas oceanográficas Mar del Plata I-V. Contribución al trazado de una carta bionómica del área de Mar del Plata. Las asociaciones del sistema litoral entre 12 y 70 m de profundidad. Boletín del Instituto de Biología Marina de Mar del Plata (Argentina) 16: 1-85.         [ Links ]

ORENSANZ JM (1973) Los anélidos poliquetos de la Provincia Biogeográfica Argentina. IV. Lumbrineridae. Physis A 32: 343-393.         [ Links ]

ORENSANZ JM (1974) Los anélidos poliquetos de la provincia biogeográfica argentina. V. Onuphidae. Physis A 33: 75-122.         [ Links ]

OYARZÚN C, FD CARRASCO & VA GALLARDO (1987) Some characteristics of macrobenthic fauna from the organic-enriched sediments at Talcahuano, Chile. Cahiers de Biologie Marine 28: 429-446.         [ Links ]

PAIVA PC (1993) Trophic structure of a shelf polychaete taxocoenosis in southern Brazil. Cahiers de Biologie Marine 35: 39-55.         [ Links ]

PEARSON TH & R ROSENBERG (1978) Macrobenthic succession in relation to organic enrichment and pollution of the marine environment. Oceanography and Marine Biology Annual Reviews 16: 229-311.         [ Links ]

PINEDO S, R SARDA & D MARTIN (1997) Comparative study of the trophic structure of soft-bottom assemblages in the Bay of Blanes (Western Mediterranean Sea). Bulletin of Marine Science 60: 529-542.         [ Links ]

POCKLINGTON P & PG WELLS (1992) Polychaetes: key taxa for marine environmental quality monitoring. Marine Pollution Bulletin 24: 593-598.         [ Links ]

PROBERT PK (1984) Disturbance, sediment stability, and trophic structure of soft-bottom communities. Journal of Marine Research 42: 893-921.         [ Links ]

REISH DJ (1955) The relation of polychaetous annelids to harbor pollution. United States Public Health Reports 70: 1168-1174.         [ Links ]

REISH DJ (1957) The relationship of the polychaetous annelid Capitella capitata (Fabricius) to waste discharges of biological origin. In: Tarzwell CM (ed) Biological problems in water pollution: 195-200. United States Public Health Service, Philadelphia, Pennsylvania.         [ Links ]

REISH DJ (1963) A quantitative study of the benthic polychaetous annelids of Bahía de San Quintin, Baja California. Pacific Naturalist 3: 399-436.         [ Links ]

REISH DJ (1980) Use of polychaetous annelids as test organisms for marine bioassay experiments. In: Buikema AL Jr & J Cairns Jr (eds) Aquatic invertebrate bioassays: 140-154. American Society for Testing and Materials, Special Technical Publication No. 715, Philadelphia, Pennsylvania.         [ Links ]

RHOADS DC & LF BOYER (1982) The effects of marine benthos on physical properties of sediments: a successional perspective. In: McCall PL & MJS Tevesz (eds) Animal-sediment relations: 3-52. Plenum Publishing Corporation, New Haven, Connecticut.         [ Links ]

ROUX A, R BASTIDA & CS BREMEC (1993) Comunidades bentónicas de la Plataforma Continental Argentina. Campañas Transección BIP "Oca Balda" 1987/88/89. Boletim do Instituto Oceanográfico de São Paulo (Brasil) 41: 81-94.         [ Links ]

RUMOHR H, E BONSDORFF & TH PEARSON (1996) Zoobenthic succession in Baltic sedimentary habitats. Archives Fisheries Marine Research 44: 179-214.         [ Links ]

RYGG B (1985) Distribution of species along pollution-induced diversity gradients in benthic communities in Norwegian fjords. Marine Pollution Bulletin 16: 469-474.         [ Links ]

SHANNON CE & W WEAVER (1963) The mathematical theory of communication. University Illinois Press, Urbana, Illinois. 117 pp.         [ Links ]

TSUTSUMI H (1990) Population persistence of Capitella sp. (Polychaeta; Capitellidae) on a mud flat subject to environmental disturbance by organic enrichment. Marine Ecology Progress Series 63: 147-156.         [ Links ]

Associate Editor: P. Camus
Received March 2, 2000; accepted September 20, 2000

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