“Amphioxus Sand” Community in Thermaikos Bay (Eastern Mediterranean)

Volume 13 – No.11a – 2004 REPRINT pp. 1122 - 1128

STRUCTURE OF THE “AMPHIOXUS SAND” COMMUNITY IN THERMAIKOS BAY (EASTERN MEDITERRANEAN)

Chryssanthi Antoniadou - Yiannis Krestenitis - Chariton Chintiroglou

Angerstr. 12 85354 Freising - Germany Phone: ++49 – (0) 8161-48420 Fax: ++49 - (0) 8161-484248 Email: [email protected] www.psp-parlar.de © by PSP Volume 13 – No 11a. 2004 Fresenius Environmental Bulletin

STRUCTURE OF THE “AMPHIOXUS SAND” COMMUNITY IN THERMAIKOS BAY (EASTERN MEDITERRANEAN)

Chryssanthi Antoniadou1, Yiannis Krestenitis2 and Chariton Chintiroglou1

1Aristotle University, School of Biology, Department of Zoology, P.O. Box 134, 540 06 Thessaloniki, Greece 2Aristotle University, School of Mechanics, Department of Hydraulics and Environmental Engineering, 540 06 Thessaloniki, Greece

SUMMARY

The structure of the “Amphioxus sand” community in expanded during the last decade as a result of increasing Thermaikos Bay was studied over a two-year period. The eutrophication and organic pollution, causing a consider- examination of the collected 4,767 specimens revealed the able decrease in B. lanceolatum’s population density and presence of 141 species. Gastropoda and Polychaeta were also a change on its community structure. Thus, this com- the dominant taxa. Multivariate analyses showed high simi- munity type seems to be threatened worldwide [19, 20]. In larity, whereas samples discriminated according to sediment the 70’s, the “Amphioxus sand” community was common granulometry. The degradation of this community was in Thermaikos Gulf, but today its presence is limited. evident in its structure as most of the recorded macrobenthic This study took place at the NE side of Thermaikos species characterizes the organically enriched sediments. Bay, where a large fishing area of the common seashell

Venus verrucosa is heavily exploited [21]. The main aim was to detect the ‘Amphioxus sand’ community, and elabo- rate its structural variability in both spatial and temporal KEYWORDS: biodiversity, soft substrate, sublittoral, Thermaikos, scales. Furthermore, the most critical factors that influence organic pollution. its distributional range were identified.

MATERIALS AND METHODS INTRODUCTION Study area Ecological studies of zoobenthos have proved to be ex- Thermaikos Gulf is a shallow-water embayment in tremely useful for biomonitoring [1-3]. Many authors have the NW Aegean (Figure 1). Its northern part, named as showed that the structure of benthic communities reflects Thessaloniki Gulf, is divided at three distinct areas: (1) the integrated conditions over a period of time [4-6]. Con- the inner bay, which is a very closed area as regards water sequently, this kind of data is helpful for pollution impact renewal, (2) the central gulf and (3) the outer gulf. Ther- studies due to anthropogenic interference [7-9]. In order to maikos receives discharges from large river systems (Ax- detect and deal with such cases, as defined by many inter- ios, Loudias, Aliakmonas) and also sewage and industrial national directives and conventions (e.g. Rio 1992, Water effluents from the city of Thessaloniki [22, 23]. The study Framework Directive, Directive 2000/60/EC), it is neces- was conducted at the NE side of the central gulf, which sary to create a database concerning the composition of covers a surface of 136 km2 with a mean depth of 25 m. natural communities [10, 11]. After preliminary sampling the “Amphioxus sand” com- Besides the intense quantitative research on soft sub- munity was found at two sites. The first site (St.1) had a strate communities, little is known about the Branchiostoma depth ranging from 3 to 5 m, whereas the second one lanceolatum facies, often called as “Amphioxus sand” [12]. (St.2), located northern, had a depth ranging from 7 to 10 m Most of the relevant information derives from studies in the (Figure 1). Western Mediterranean and the Adriatic [13-17], as there is only one reference from the Eastern basin [18] and the Black Sampling techniques Sea [19]. The“Amphioxus sand” community normally ex- Abiotic Factors: The main abiotic factors, e.g. tempera- tends on gravel and coarse sand with shell fragments, often ture, salinity, dissolved O2 and pH, were measured in the accompanied by certain Polychaeta species (e.g. Stauro- water column with a CTD on a monthly basis. Currents were cephalus keffersteini) and venerupid bivalves (e.g. Venerupis recorded with an autographic current-meter (AANDERAA aurea, Venus verrucosa) [12]. However, muddy areas have RCM9) placed five meters beyond see level. A sediment

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FIGURE 1 - Map of the study area indicating sampling sites.

analysis (granulometric composition and amount of organic The data obtained per sampling site were analyzed material contained in sediment) was carried out at each with multidimensional scaling techniques, based on the sampling site on a seasonal basis. Two substrate samples Bray-Curtis similarity and log transformed numerical were collected with a core sampler of 1 lt volume, at each abundances, using PRIMER package [30]. The signifi- period with SCUBA diving. The organic material was cance of the multivariate results was assessed with measured with the H2O2 method and the identification of ANOSIM test. SIMPER analysis was applied to identify the granulometric composition was based on Folks system the contribution of each species to the overall similarity of sediment classification [24, 25]. within a group of samples and the dissimilarity among groups [30]. Finally, the BIOENV procedure was used to Macrofauna examine which environmental parameters are related to Sampling was carried out in August and January for the observed biotic pattern (MDS plot) and the degree of two consecutive years (summer 2001 to winter 2003). Mac- this relation [30]. rofaunal samples were collected from each site and period by SCUBA diving with the use of an 18 x 25 cm core sampler (3 replicates from the shallow-depth site and 5 repli- RESULTS cates from the deeper one) [26, 27]. Overall, 32 samples were obtained, sieved (mesh opening 0.5 mm) and fixed Community structure in 9% formaldehyde. After sorted, all living specimens A total of 4,767 individuals were counted, belonging to were identified at species level and counted. 142 species. From these species 55 were dominant (15 Poly- chaeta, 36 Mollusca, 4 Crustacea), according to population Data analysis density values and frequency (> 60%) (Table 1). Data was analyzed by standard biocoenotic methods. Gastropods dominate in the first year and polychaetes Thus, the population density, the mean dominance, the in the second. MANOVA results showed that from the frequency, and also Margalef’s richness, Shannon-Wiener four dominant taxa only Gastropoda had unequal disper- and Pielou’s evenness indices, were calculated [27, 28]. sion of abundance in time and Bivalvia in both time and Numerical abundance data were analyzed by mANOVA, space (Table 2). in order to examine the effect of three different factors: a) The values of diversity indices were high (Figure 2). site, b) season and c) year of sampling. A logarithmic trans- Richness values (d) ranged from 7.47 to 9.77, H’ values formation (logx+1) was used to normalize the variance of from 3.42 to 4.43 and J’ values from 0.72 to 0.95, indicat- numerical abundances [29]. ing the evenly dispersion of numerical abundance among species.

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TABLE 1 - Taxonomic list of the recorded species.

Taxa - Species Cnidaria Bittium reticulatum (DaCosta, 1778) Lucinella divaricata (Linnaeus, 1758) Cereus pedunculatus (Pennant, 1777) Bolinus brandaris (Linnaeus, 1758) Macoma cumana (Costa O.G., 1829) Polychaeta Caecum trachea (Montagu, 1803) Mactra corallina (Linnaeus, 1758) Abarenicola claparedei (Levinsen, 1883) Cerithiopsis horrida (Monterosato, 1874) Modiolus barbatus (Linnaeus, 1758) Aonides oxycephala (Sars, 1862) Cerithium vulgatum Bruguiere, 1792 Mysia undata (Pennant, 1777) Aphelochaeta marioni Saint-Josephl, 1894 Chrysallida doliolum (Philippi, 1844) Mytilus galloprovincialis Lamarck, 1819 Aponuphis bilineata (Baird, 1870) Clanculus jussieui (Payraudeau, 1826) Nucula nitidosa Winckworth, 1930 Aricidea fragilis mediterranea Laubier & Ramos, Conus mediterraneus Hwass, in Bruguiere, 1792 Nucula sulcata Bronn, 1831 1974 Branchiomma bombyx (Dalyell, 1853) Cytharella coarctata (Forbes, 1840) Nuculana pella (Linnaeus, 1767) Capitella capitata (Fabricius, 1780) Epitonium sp. Paphia aurea (Gmelin, 1791) Caulleriella bioculata (Keferstein, 1862) Eulimella scillae (Scacchi, 1835) Parvicardium exiguum (Gmelin, 1791) Ceratonereis costae (Grube, 1840) Euspira catena (DaCosta, 1778) Pita rudis (Poli, 1795) Chone filicaudata Southern, 1914 Folinella excavata (Philippi, 1836) Psammobia depressa (Pennant, 1777) Cirriformia tentaculata (Montagu, 1808) Fusinus rudis (Philippi, 1844) Scrobicularia plana (DaCosta, 1778) Diopatra neapolitana DelleChiaje, 1841 Fusinus syracusanus (Linnaeus, 1758) Tellina donacina (Linnaeus, 1758) Eteone picta Quatrefages, 1865 Gibbula adansonii (Payraudeau, 1826) Tellina planata (Linnaeus, 1758) Eunice vittata (DelleChiaje, 1829) Hadriana oretea (Brocchi, 1814) Tellina pulchella Lamarck, 1818 Euclymene oerstedi (Claparede, 1863) Haminaea navicula (DaCosta, 1778) Venus verrucosa (Linnaeus, 1758) Exogone naidina Oersted, 1845 Hexaplex trunculus (Linnaeus, 1758) Cirripedia Glycera tridactyla Schmarda, 1861 Jujubinus striatus (Linnaeus, 1758) Balanus trigonus Darwin, 1854 Grubeosyllis limbata (Claparede, 1868) Leiostraca subulata (Donovan, 1804) Amphipoda Hydroides elegans (Haswell, 1883) Mangelia attenuata (Montagu, 1803) Ampelisca diadema (Costa, 1853) Levinseria grasilis (Tauber, 1879) Manzonia crassa (Kanmacher, 1798) Amphithoe ramondi Audouin, 1826 Lumbrineris latreilli Audouin & Milne-Edwards, Monophorus perversus (Linnaeus, 1758) Caprella acanthifera Leach, 1814 1834 Maldane glebifex Grube, 1860 Nassarius incrassatus (Strom, 1768) Corophium sextonae Crawford, 1937 Melinna palmata Grube, 1870 Nassarius limata (Chemnitz, 1795) Dexamine spinosa (Montagu, 1813) Marphysa bellii (Audouin & Milne-Edwards, 1833) Nassarius reticulatus (Linnaeus, 1758) Elasmopus brasiliensis (Dana, 1855) Naineris laevigata (Grube, 1855) Neverita josephinia Risso, 1826 Elasmopus rapax Costa, 1853 Nematonereis unicornis (Grube, 1840) Odostomia conoidea (Brocchi, 1814) Ericthonius brasiliensis (Dana, 1855) Nephtys incisa Malmgren, 1865 Pusillina radiata (Philippi, 1836) Microdeutopus anomalus (Rathke, 1843) Nereiphylla rubiginosa (Saint-Joseph, 1888) Ringicula auriculata (Menard de la Groye, 1811) Perioculodes aequimanus (Korssman, 1880) Notomastus latericeus Sars, 1851 Rissoa guerinii Recluz, 1843 Phtisica marina Slabber, 1749 Notomastus profundus Eisig, 1887 Rissoa monodonta Philippi, 1836 Tanaidacea Paralacydonia paradoxa Fauvel, 1913 Rissoa violacea Desmarest, 1814 Leptochelia savignyi (Kroyer, 1842) Phyllodoce mucosa Oersted, 1843 Smaragdia viridis (Linnaeus, 1758) Isopoda Pista cristata (O.F. Muller, 1776) Tricolia pullus (Linnaeus, 1758) Astacilla longicornis (Sowerby, 1806) Piromis eruca (Claparede, 1870) Trivia sp. Dynamene bidendatus (Adams, 1800) Protodorvillea kefersteini (McIntosh, 1869) Trophon muricatus (Montagu, 1803) Paranthura nigropunctata (Lucas, 1849) Polyopthalmus pictus (Dujardin, 1839) Turbonilla lactea (Linnaeus, 1758) Sphaeroma serratum (Fabricius, 1787) Serpula vermicularis Linnaeus, 1767 Turbonilla rufa (Philippi, 1836) Cumacea Syllidia armata Quatrefages, 1865 Turritella communis Risso, 1826 Cumela limicola Sars, 1879 Syllis hyalina Grube, 1863 Vexillum tricolor (Gmelin, 1791) Decapoda Syllis prolifera Krohn, 1852 Scaphopoda Cestopagurus timidus (Roux, 1830) Sipuncula Dentalium dentale Linnaeus, 1767 Pisidia longimana (Risso, 1816) Phascolosoma granulatum Leuckart, 1828 Bivalvia Thoralus cranchii (Leach, 1817) Nemertina Abra alba (Wood W., 1802) Callianassidae Polyplacophora Acanthocardia echinata (Linnaeus, 1758) Echinoidae Lepidochitona scabridus (Jeffreys, 1880) Chlamys glabra (Linnaeus, 1758) Spatangus sp. Gastropoda Corbula gibba (Olivi, 1792) Ophiuroidae Ophiothrix fragilis (Abildgaard in Muller, Acteon tornatilis (Linnaeus, 1758) Donax variegatus Gmelin, 1790 1789) Alvania cimex (Linnaeus, 1758) Dosinia lupinus (Linnaeus, 1758) Olothuroidae Alvania discors (Allan, 1818) Gastrana fragilis (Linnaeus, 1758) Olothuria polii DelleChiaje, 1823 Bela ginaria (Risso, 1826) Glans trapezia (Linnaeus, 1767) Branchiostoma lanceolatum (Pallas, 1774) * indicate dominant species

TABLE 2 - MANOVA results.

Site Season Year Taxa F p F p F p Polychaeta 0.03 0.22 0.04 0.83 0.19 0.66 Gastropoda 1.56 0.22 0.21 0.64 30.8 0.00 Bivalvia 11.08 0.00 11.13 0.00 0.12 0.73 Crustacea 0.37 0.55 0.69 0.41 0.11 0.74 Significant differences in bold

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80 1000 Ν umerical abundance minor annual grouping is apparent, whereas inside group C a seasonal pattern during the second year came up. The 60 800 600 discrimination of the above pattern is confirmed by one- 40 way ANOSIM (global R = 0.81; p < 0.1). The pairwise 400 test showed significant differences among all group com- 20 200

Species richness Species binations (R ranging from 0.79 to 1). The SIMPER analy- 0 0 sis identified 18 species as most contributing to in-group St.1 St.2 St.1 St.2 St.1 St.2 St.1 St.2 similarity and 45 species to among-groups dissimilarity Summer 2001 Winter 2002 Summer 2002 Winter 2003 (Table 3).

d H' J' S01 12 1 W02 A W02 B ' 10 0,8 Index J' W03 Η S01 8 0,6 St.1 6 S02 0,4 4 St.2

Indices d, Indices 2 0,2 W03 C 0 0 S02 St.1 St.2 St.1 St.2 St.1 St.2 St.1 St.2 stress = 0.02 Summer 2001 Winter 2002 Summer 2002 Winter 2003 FIGURE 3 - Non-metric multidimensional scaling, FIGURE 2 - Temporal and spatial variation of based on Bray-Curtis similarity index, calculated diversity indices (d = Margalef’s richness, H’ = from root transformed numerical abundance data Shannon-Wiener index, J’ = Pielou’s evenness). The BIOENV procedure showed that the substrates Affinity of sites granulometry, and especially the proportion of gravels Non-metric MDS indicates the separation of the sam- (SR = 0.56) and of sand (SR = 0.35) are the factors that ples in three main groups according to sampling site and relate mostly with the community structure. the year of sampling (Figure 3). Inside group A, a second

TABLE 3 - Species contributing on about 50% of the average in-group similarity or the average among groups dissimilarity.

A B C A-B A-C B-C Taxa 42.3 55.1 60.2 59.1 60.7 52.8 Abra alba 4.06 1.61 2.49 Ampelisca diadema 3.09 2.19 1.59 1.08 Aonides oxycephala 5.91 1.95 3.12 2.52 Aricidea fragilis 1.49 1.46 Bittium reticulatum 6.80 10.33 8.77 2.08 2.00 1.39 Caecum trachea 4.06 1.99 2.57 Capitella capitata 11.62 7.32 8.28 Cauleriella bioculata 1.36 1.36 Cestopagurus timidus Chone filicaudata 5.07 4.80 1.92 1.37 Chrysallida doliolum 1.47 1.90 Clanculus jussieui 1.64 1.71 Corbula gibba 1.50 Corophium sextonae 1.14 Cytharella coarctata 6.19 2.39 1.21 3.76 Dentalium dentale 5.51 3.08 3.09 Elasmopus rapax 1.96 2.09 Euspira catena 1.58 1.97 Gibbula adansonii 4.87 2.25 2.82 Glycera tridactyla 5.07 3.09 Hadriana oretea 1.10 1.31 Lucinella divaricata 1.16 1.28 1.15 Lumbrineris latreilli 4.00 Macoma cumana 1.22 Maldane glebifex 2.03 1.37 Manzonia crassa 1.18 Melinna palmata 1.25 1.94 1.54

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TABLE 3 (continued).

A B C A-B A-C B-C Taxa 42.3 55.1 60.2 59.1 60.7 52.8 Modiolus barbatus 1.29 1.97 Mysia undata 1.35 1.60 Nassarius incrassatus 4.58 1.09 1.64 Nassarius limata 4.22 2.30 2.90 Neverita josephinia 1.41 1.31 Notomastus latericeus 1.29 1.31 Nucula sulcata 1.59 1.44 Odostomia conoidea 1.32 1.32 Paranthura nigropunctata 1.36 Piromis eruca 1.30 1.38 Pisidia longimana 1.47 Pista cristata 1.47 1.51 Protodorrillea kefersteini 6.21 2.83 2.03 1.16 Pusillina radiata 9.61 8.14 6.72 1.23 1.27 Ringicula auriculata 1.34 Smaragdia viridis 5.04 1.39 2.01 Syllis prolifera 1.19 1.25 Tellina planata 1.30 1.22 Tellina pulchella 1.35 1.20 Tricolia pullus 5.04 2.13 2.64 Trivia sp. 1.54 1.63 Turbonilla lactea 1.39 Total 54.2 51.4 51.9 50.2 50.9 50.1

DISCUSSION only at St.2 in the second year, is considered as an indica- tion of deterioration of benthic temperate communities [6]. The Mediterranean coastal zone has experienced The separation of the two sites is also attributed on their rapid urban development and industrialization in recent granulometric differences. Thus, St.2 contains a higher years [3, 31]. Thermaikos gulf is considered among the proportion of shell fragments and silt that diversifies its most anthropogenically disturbed marine area in Greece, macrobenthic fauna. The organic load in sediment was since it receives both domestic and industrial waste, from found increased at both sites, over-passing 1-2 %. As the the city of Thessaloniki and its adjacent industrial area differences on organic amount were insignificant among [22, 32]. The management of this gulf is difficult; since on sites, periods and years, this factor was not related with one hand, the estuary system of the rivers Axios Loudias, the biotic pattern. Gallikos and Aliakmon is under protection by the Ramsar convention, and, on the other there is an extended aqua- The basic concept of biodiversity focuses on species culture of the common Mediterranean mussel [32]. So, the richness, e.g. composition that is the important indicator entire gulf is strongly affected by human activities. Re- of diversity across spatial scales and habitats [10, 11]. cent research efforts focus on the assessment of pollution Comparing our results with relevant descriptions of the effects on marine benthos, by investigating the response “Amphioxus sand” community we can note its diversified of benthic communities to different disturbance factors [6, composition (Table 4). The greater number of macroben- 28, 33]. thic species was recorded in 1970 in the Western Mediter- ranean, whereas the lower biodiversity was reported from The majority of relevant data concerning soft substra- the Adriatic and the Black Sea. tum in Thermaikos gulf, derive from NCMR technical reports [34]. However, Patoucheas and Stamou [35] de- The species richness in the present study has a rather veloped a multivariate Markovian model and concluded high value. Regarding the ecological preferences of the that dissolved oxygen is the main factor, which controls recorded species, the relevant information is limited on the structure of benthic communities in Thermaikos gulf. polychaetes and peracarids. Thus, many of the dominant Our results showed that the current biotic pattern is less species are characterized as tolerant or related to organic affected by dissolved oxygen than by the sediment clusters. pollution, constituted the 65% of the recorded polychaete It is known that Branchiostoma lanceolatum, an inhabitant species (i.e. Aphelochaeta marioni, Caulleriella bioculata, of the sandy sublittoral zone, is favored by the presence of Capitella capitata, Chone filicaudata, Cirriformia tentacu- fragmented shells and gravels [16, 19]. Thus, the change lata, Eteone picta, Εunice vittata, Exogone naidina, Eucly- to the proportions of sediment compounds among the two mene oerstedi, Glycera tridactyla, Hydroides elegans, Levin- years of survey had a strong impact on the structure of the senia gracilis, Lumbrineris latreilli, Maldane glebifex, community, resulting to an annual discrimination. The loss Melinna palmata, Marphysa bellii, Nepthys incisa, Notomas- of seasonality in community structure, which reappears tus latericeus, N. profundus, Paralacydonia paradoxa,

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TABLE 4 - Composition of the “Amphioxus sand” community.

Guille 1970 Dounas 1986 Rellini 1986 Konsulova 1991 Taxa Present study Marseille Strymonikos Gulf Cinque Terre Varna Bay Porifera 3 3 Cnidaria 9 1 Polychaeta 53 55 5 28 40 Nemertina 1 1 Mollusca 34 22 12 15 74 Crustacea 47 64 3 16 22 Tunicata 3 1 Echinodermata 25 9 3 3 Sipuncula 2 3 1 1 Pycnogonida 1 Bryozoa 5 Total 183 160 25 61 141

Phyllodoce mucosa, Pista cristata, Piromis eruca, Proto- [4] Bellan, G., Bellan-Santini, D. and Picard, J. (1978). Les mo- dorvillea keffersteini, Syllis hyalina and Syllis prolifera) dalités de répartition, en Méditerranée nord-orientale, des pe- uplements benthiques des sédiments côtiers soumis a la pol- and the 45% of amphipods (i.e. Ampelisca diadema, Coro- lution par matières organiques dominantes. IV Journées Etud. phium sextonae, Elasmopus brasiliensis, E. rapax, Erich- Pollutions, Antalaya, C.I.E.S.M., : 351-358. thonius brasiliensis), whereas the only recorded Tanaidacea (i.e. Leptochelia savignyi) is favoured by organic load [3, [5] Gray, J. (1979). Pollution-induced changes in populations. 33, 36-38]. Philosoph. Trans. Royal Soc. London, SeriesB : Biol. Sci., 228: 545-561. Muddy areas have expanded during the last decade as a result of increased eutrophication and organic pollution, [6] Karalis, P., Antoniadou, C. and Chintiroglou, C. (2003). Structure of the artificial hard substrate assemblages in ports causing a severe decline of Branchiostoma lanceolatum in North Aegean Sea (Thermaikos Gulf). Oceanologica Acta, populations on global scale [12, 19, 20, 31]. In the 60’s the 26: 215-224. species was quite common along the Mediterranean coasts but today it is considered rare [19]. In Thermaikos Gulf [7] Grant, A. and Briggs, A. (2002). Toxicity of sediments from Β. lanceolatum was recorded during the ΜEDPOL expedi- around a North Sea oil platform: are metals or hydrocarbons responsible for ecological impacts? Mar. Envir. Res., 53 : 95- tion (1980), whereas no quantitative data on its population 116. density or its community structure have been reported. Today the “Amphioxus sand” community appears in the [8] Nicolaidou, A., Pancucci, M.A. and Zenetos, A. (1989). The NE area, however, its community structure is altered, impact of dumping coarse metalliferouw waste on the ben- since most of the dominant species are characteristics of thos in Evoikos Gulf, Greece. Mar. Poll. Bull., 20(1) : 28-33 the organically polluted sediments. Concluding, the effect of the organic pollution, well reported for the broader area [9] Lu, L. and Wu, R.S.S. (2003). Recolonization and succession of subtidal macrobenthic infauna in sediments contaminated of Thessaloniki Bay [22] and evaluated from the organic with cadmium. Environ. Poll., 121 : 27-38. compound in our sediment analysis, was found to be rather indirect than direct on the current biotic pattern, as [10] Gaston, K. and Spicer, J.I. (1996). Biodiversity. An introduc- the higher proportion of species was ecologically classi- tion. Blackwell, Oxford, 1-113. fied among the indicators of the organically enriched sediments. [11] Costello, M.J. (1998). To know, research, manage and con- serve marine biodiversity. Océanis, 24: 25-49.

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